Content uploaded by Shi-Ke Huang
Author content
All content in this area was uploaded by Shi-Ke Huang on Nov 11, 2021
Content may be subject to copyright.
Vol.:(0123456789)
1 3
Fungal Diversity
https://doi.org/10.1007/s13225-021-00488-4
Taxonomic studies ofsome often over‑looked Diaporthomycetidae
andSordariomycetidae
Shi‑KeHuang1,2,3,4,5· KevinD.Hyde3,6· AusanaMapook3· SajeewaS.N.Maharachchikumbura7·
Jayarama D.Bhat8,9 · EricH.C.McKenzie10· RajeshJeewon11· Ting‑ChiWen1,2,5
Received: 11 May 2021 / Accepted: 10 August 2021
© MUSHROOM RESEARCH FOUNDATION 2021
Abstract
Sordariomycetes is an earlier-introduced and one of the widely distributed class of Ascomycota. The class was initially clas-
sified based on morphology in having inoperculate and unitunicate asci. With the development of DNA based phylogenetic
analysis, several undetermined or polyphyletic members of Sordariomycetes were reclassified. However, not all species
belonging to this class have been sequenced and analyzed. There are a number of species, especially those old and poorly
studied ones which have never been sequenced before and not even recollected again for further taxonomic verification. One
of the main objective in this study is to revise and update the taxonomy of several well-known early and poorly studied spe-
cies whose classification are still obscure. Herein, we re-examined the type materials and/or authentic specimens together
to explore 74 relatively poorly-studied genera, which mainly belong to Boliniales, Calosphaeriales, Chaetosphaeriales,
Jobellisiales, and Sordariales classified under Diaporthomycetidae and Sordariomycetidae. We provide descriptions, notes,
figures and/or drawings and discussed their phylogenetic relationships. As a result, the monotypic Jobellisiales is transferred
from Hypocreomycetidae to Diaporthomycetidae. Based on phylogenetic analysis, the polyphyletic Lasiosphaeriaceae is
divided into five families, Bombardiaceae (Apodospora, Bombardia, Bombardioidea, Fimetariella and Ramophialophora),
Lasiosphaeriaceae (Anopodium, Bellojisia, Corylomyces, Lasiosphaeria, Mammaria and Zopfiella), Lasiosphaeridaceae
(Lasiosphaeris), Strattoniaceae (Strattonia) and Zygospermellaceae (Episternus and Zygospermella). In addition, a new
family Neoschizotheciaceae is established based on Neoschizothecium. Analysis of the type species of Boothiella, Stel-
latospora, Sulcatistroma and Tengiomyces placed them in Sordariaceae, Chaetomiaceae, Hypocreales and Coronophorales,
respectively. We classify the genera lacking molecular data based on their morphology and expect them to be recollected; that
is, Kacosphaeria in Calosphaeriales; Arnium, Biconiosporella, Camptosphaeria, Diffractella, Emblemospora, Eosphaeria,
Periamphispora, Synaptospora and Tripterosporella in Sordariales; Conidiotheca in Sordariomycetes; Copromyces, Effetia,
Endophragmiella and Tulipispora are accommodated in Ascomycota. Besides, we establish a new genus Neoschizothecium
based on phylogenetic analysis. New combinations proposed: Camaropella amorpha, Cam. microspora, Cam. plana, Clad-
orrhinum grandiusculum, Cla. leucotrichum, Cla. terricola, Cla. olerum, Helminthosphaeria plumbea, Immersiella hirta,
Jugulospora minor, Lasiosphaeris arenicola, Neoschizothecium aloides, Neo. carpinicola, Neo. conicum, Neo. curvisporum,
Neo. fimbriatum, Neo. glutinans, Neo. inaequale, Neo. minicaudum, Neo. selenosporum, Neo. tetrasporum, Neurospora
autosteira, Podospora brunnescens, P. flexuosa, P. jamaicensis, P. hamata, P. macrospora, P. spinosa, Strattonia petrogale
and Triangularia microsclerotigena, T. nannopodalis, T. praecox, T. samala, T. tarvisina, T. unicaudata, T. yaeyamensis.
New epithets are proposed for Apiorhynchostoma apiosporum and Podospora dacryoidea.
Keywords 43 new taxa· Boliniales· Calosphaeriales· Helminthosphaeriaceae· Jobellisiales· Sordariales·
Sordariomycetes
Handling Editor: Jian-Kui Liu.
* Ting-Chi Wen
tingchiwen@yahoo.com
Extended author information available on the last page of the article
Fungal Diversity
1 3
Table ofcontents
Arrangement of genera in Boliniaceae, Calospha-
eriaceae, Helminthosphaeriaceae, Jobellisiaceae, Lasio-
sphaeriaceae, Pleurostomataceae, Podosporaceae and
Sordariaceae. Those not in agreement with Wijayawardene
etal. (2020) are marked with an asterisk.
Phylum Ascomycota Caval.-Sm.
Class Sordariomycetes O.E. Erikss. & Winka
Subclass Diaporthomycetidae Senan., Maharachch. &
K.D. Hyde
Calosphaeriales M.E. Barr
Calosphaeriaceae Munk
Calosphaeria Tul. & C Tul.
Flabellascus Réblová
Jattaea Berl.
Togniniella Réblová etal.
Pleurostomataceae Réblová etal.
Pleurostoma Tul. & C. Tul
Calosphaeriales genus incertae sedis
Kacosphaeria Speg.
Jobellisiales M.J. D’souza & K.D. Hyde*
Jobellisiaceae Réblová
Jobellisia M.E. Barr
Other genera in Diaporthomycetidae
Ophiostomatales Benny & Kimbr.
Ophiostomataceae Nannf.
Fragosphaeria Shear
Subclass Hypocreomycetidae O.E. Erikss. & Winka
Genera transferred to Hypocreomycetidae
Coronophorales genus incertae sedis
Tengiomyces Réblová*
Hypocreales genus incertae sedis
Sulcatistroma A.W. Ramaley*
Subclass Sordariomycetidae O.E. Erikss. & Winka
Boliniales P.F. Cannon
Boliniaceae Rick
Apiocamarops Samuels & J.D. Rogers
Apiorhynchostoma Petr.
Camaropella Lar.N. Vassiljeva
Camarops P. Karst.
= Bolinia (Nitschke) Sacc.
Cornipulvina Huhndorf etal.
Endoxyla Fuckel
Mollicamarops Lar.N. Vassiljeva
Neohypodiscus J.D. Rogers etal.
Pseudovalsaria Spooner
Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández
Helminthosphaeriaceae Samuels, Cand. & Magni
Echinosphaeria A.N. Mill. & Huhndorf
Helminthosphaeria Fuckel
Hilberina Huhndorf & A.N. Mill.
Ruzenia O. Hilber
Sordariales Chadef. ex D. Hawksw. & O.E. Erikss
Bombardiaceae S.K. Huang & K.D. Hyde*
Apodospora Cain & J.H. Mirza*
Bombardia (Fr.) P. Karst.*
Bombardioidea C. Moreau ex N. Lundqv.*
Fimetariella N. Lundq.*
Ramophialophora M. Calduch etal.*
Lasiosphaeriaceae Nannf.
Anopodium N. Lundq.
Bellojisia Réblová
Corylomyces Stchigel etal.
Lasiosphaeria Ces. & De Not.
Mammaria Ces. ex Rabenh.
Zopfiella G. Winter
Lasiosphaeridaceae S.K. Huang & K.D. Hyde*
Lasiosphaeris Clem.*
Podosporaceae X. Wei Wang & Houbraken
Cladorrhinum Sacc. & Marchal
Podospora Ces.
= Schizothecium Corda*
Triangularia Boedijn
= Apiosordaria Arx & W. Gams*
Sordariaceae G. Winter
Boothiella Lodhi & Mirza*
Guilliermondia Boud.
Neurospora Shear & B.O. Dodge
Pseudoneurospora Dania García etal.
Sordaria Ces & De Not.
Strattoniaceae S.K. Huang & K.D. Hyde*
Strattonia Cif.*
Zygospermellaceae S.K. Huang & K.D. Hyde*
Episternus Górz & Boroń*
Zygospermella Cain*
Fungal Diversity
1 3
Other genera in Sordariales
Chaetomiaceae G. Winter
Stellatospora T. Ito & Nakagiri*
Neoschizotheciaceae S.K. Huang & K.D. Hyde*
Apodus Malloch & Cain*
Cercophora Fuckel*
Echria (N. Lundq.) Kruys etal.*
Immersiella A.N. Mill. & Huhndorf*
Jugulospora N. Lundq.*
Neoschizothecium S.K. Huang & K.D. Hyde*
Rinaldiella Deanna A. Sutton etal.*
Zygopleurage Boedijn*
Sordariales genera incertae sedis
Arnium Nitschke ex G. Winter*
Biconiosporella Schaumann*
Camptosphaeria Fuckel*
Diffractella Guarro etal.*
Emblemospora Jeng & J.C. Krug*
Eosphaeria Höhn.*
Isia D. Hawksw. & Manohar.
Lockerbia K.D. Hyde
Periamphispora J.C. Krug*
Reconditella Matzer & Hafellner
Roselliniopsis Matzer & Hafellner
Synaptospora Cain*
Tripterosporella Subram. & Lodha*
Other genera studied
Subclass Xylariomycetidae O.E. Erikss. & Winka
Xylariales Nannf.
Diatrypaceae Nitschke
Monosporascus Pollack & Uecker
Sordariomycetes genus incertae sedis
Conidiotheca Réblová & L Mostert *
Ascomycota genera incertae sedis
Copromyces N. Lundq.*
Effetia Bartoli etal.*
Endophragmiella B. Sutton*
Tulipispora Révay & J. Gönczöl *
Introduction
The class Sordariomycetes was introduced by Eriksson
and Winka (1997) and is classified under the subdivision
Pezizomycotina (Hyde etal. 2013, 2020; Maharachchikum-
bura etal. 2015, 2016; Wijayawardene etal. 2018, 2020).
The numbers of orders, families and genera accepted in
Sordariomycetes has been debated, with authors accepting
different numbers. Kirk etal. (2008) included 15 orders, 64
families and 1119 genera, Lumbsch and Huhndorf (2010)
included 18 orders, 63 families and 947 genera, Maharach-
chikumbura etal. (2015) included 32 orders, 105 families
and 1331 genera and Wijayawardene etal. (2018) included
37 orders, 134 families and 1060 genera. In the most recent
treatments of Sordariomycetes, seven subclasses, 45 orders,
167 families and 1506 genera were accepted with more than
20,000 species (Hyde etal. 2020), while Wijayawardene
etal. (2020) accepted 50 orders, 188 families and 1413
genera. Species in this class are widely distributed and
include plant pathogenic (Jayawardena etal. 2019), endo-
phytic (Rashmi etal. 2019), saprobic (Luo etal. 2019),
epiphytic (Hongsanan etal. 2016) and coprophilous (Kruys
etal. 2014), fungicolous (Sun etal. 2019) and lichenicol-
ous (Muggia etal. 2015) taxa. They are found in terrestrial,
aquatic and marine habitats (Hyde and Wong 2000; Cai
etal. 2006b; Jones etal. 2015, 2020; Maharachchikumbura
etal. 2016; Luo etal. 2019; Bundhun etal. 2020; Hyde etal.
2020, 2021).
Traditionally, taxa in Sordariomycetes were characterized by
inoperculate, unitunicate asci. However, some members were
reported with fissitunicate asci (the inner and the outer ascus
wall layer almost completely detach from each other with the
endoascus extruding out of the exoascus when ascus dehiscence)
(Bellemère 1994; Damm etal. 2008; Maharachchikumbura etal.
2016; Hyde etal. 2020). Traditional morphology helped classify
and understand species, however, phenotypes largely overlap
and fluctuate under different environmental conditions (Trail
etal. 2017). Therefore, combining DNA sequence analyses with
morphological studies to determine the affinities and taxonomic
placement of Sordariomycetes and other species have been very
useful to better understand their classification. Initially, phylo-
genetic analyses for Sordariomycetes were based on the internal
transcribed spacer (ITS), nuclear ribosomal large subunit (LSU)
and nuclear ribosomal small subunit (SSU) sequence data (Ber-
bee and Taylor 1992; Spatafora and Blackwell 1993; Spatafora
1995). Later protein coding gene regions (e.g. translation elon-
gation factor (TEF), beta-tubulin (TUB) and RNA polymerase II
(RPB2)) were used to provide a more stable classification of this
group (Zhang etal. 2006; Tang etal. 2007). Currently, rDNA
sequences data combined with protein coding genes are widely
recommended to resolve taxonomic placements in Sordariomy-
cetes (Maharachchikumbura etal. 2015; Hyde etal. 2017, 2020;
Hongsanan etal. 2017). Hyde etal. (2017, 2020) analyzed DNA
sequence data with divergent time estimates to investigate evo-
lutionary processes in Sordariomycetes.
Sordariomycetidae, as the earliest subclass of Sordariomy-
cetes, accommodated eight orders based on molecular data and
usually has dark ascomata, periphysate ostioles and unituni-
cate asci (Eriksson and Winka 1997; Maharachchikumbura
etal. 2016; Hyde etal. 2020). The members of this subclass
Fungal Diversity
1 3
are widely distributed on various substrates worldwide and
historically, the classification of these taxa were mainly deter-
mined using morphology and host (Eriksson and Winka 1997;
Maharachchikumbura etal. 2015, 2016; Wang etal. 2019a,
b; Hyde etal. 2020; Marin-Felix etal. 2020). Especially for
Sordariales, which was established by Hawksworth and Eriks-
son (1986) and most species have membraneous or coriaceous
ascomata and hyaline or brown, cylindrical or ellipsoidal or
irregular-shaped ascospores often with appendages or a sheath
and grow on dung or decaying substrates (Lundqvist 1972;
Zhang etal. 2006; Maharachchikumbura etal. 2015, 2016).
Huhndorf etal. (2004b) divided this order into three families,
Chaetomiaceae, Lasiosphaeriaceae and Sordariaceae, based
on morphology and phylogenetic analyses. Traditionally, a
large number of Lasiosphaeriaceae species grow on decayed
substrates and have membraneous or coriaceous ascomata
and ellipsoidal to cylindrical ascospores, usually with append-
ages (Lundqvist 1972; Hilber and Hilber 1979; Shearer 1989;
Guarro etal. 1991, 1996; Bell 2000). Most species that grow
on faeces and characterized by dark ascomata and ellipsoidal
or irregular ascospores, sometimes with ribs or pits on the
surface or surrounded by gelatinous sheath, were included in
Sordariaceae (Cain and Mirza 1969; Lundqvist 1972; Jeng
and Krug 1976; Abdullah and Rattan 1978; Hawksworth and
Manoharachary 1978; Krug 1989; Ito and Nakagiri 1994). Sub-
sequently, traditionally defined genera assumed to be in Lasio-
sphaeriaceae and Sordariaceae were either found to be polyphy-
letic or found not to be close relative of these families based on
DNA sequence analyses (García etal. 2004; Cai etal. 2006a,
b; Huhndorf etal. 2004b; Nygren etal. 2011; Kruys etal. 2014;
Wang etal. 2019a). Thus, several lasiosphaeriaceous taxa were
reassigned to establish Diplogelasinosporaceae, Naviculispo-
raceae, Podosporaceae and Schizotheciaceae, and the remain-
ing species were placed in Lasiosphaeriaceae sensu lato (Wang
etal. 2019a; Marin-Felix etal. 2020). Schizotheciaceae was
established based on Schizothecium, which is a synonym of
Podospora (Podosporaceae) (Wang etal. 2019a; Marin-Felix
etal. 2020; Index Fungorum 2020). Stellatospora was trans-
ferred from Sordariaceae to Chaetomiaceae based on available
molecular data analysis (Wang etal. 2019a). Moreover, Neuros-
pora has three mating strategies and ascospores with ribs or pits
on surface (García etal. 2004; Cai etal. 2006b); Sordaria has
smooth-walled ascospores surrounded by a gelatinous sheath
or sheath absent (Moreau 1953; Lundqvist 1972; von Arx etal.
1987; Cai etal. 2006b), but both genera are polyphyletic and
these morphologies are not consistent with the classification
recovered from DNA based phylogenies (García etal. 2004; Cai
etal. 2006b; this study). In addition, Wang etal. (2016, 2019a,
2019b) redefined several large genera in Chaetomiaceae, such
as Chaetomium, Humicola and Thielavia, and re-circumscrip-
tion of these taxa following phylogenetic analyses.
The monotypic Boliniales is widespread and usually
saprobic on wood, and it was initially characterized by
membranaceous ascomata, cylindrical asci and ellipsoidal
ascospores (Kirk etal. 2001). Subsequently, several species
with carbonaceous ascomata were classified into this order
(Zhang etal. 2006). Thus, the monophyletic Boliniales was
reported to form sister relationships to Phyllachorales in Sor-
dariomycetidae and has cylindrical asci and smooth-walled
ellipsoidal ascospores with germ pore(s) (Maharachchikum-
bura etal. 2016; Hongsanan etal. 2017; Hyde etal. 2017;
this study).
Chaetosphaeriaceae, Helminthosphaeriaceae, Leptosporel-
laceae and Linocarpaceae were accommodated in Chaeto-
sphaeriales based on molecular data analysis, and they are
widely distributed as saprobic, pathogenic or fungicolous
fungi (Huhndorf etal. 2004a; Maharachchikumbura etal.
2016; Hernández-Restrepo etal. 2017; Konta etal. 2017;
Hyde etal. 2020). As the first introduced family in this order,
Helminthosphaeriaceae contained seven genera, Echinospha-
eria, Endophragmiella, Helminthosphaeria, Hilberina, Ruze-
nia, Synaptospora and Tengiomyces, which are characterized
by hairy ascomata, cylindrical to clavate asci and cylindrical
ascospores (Miller and Huhndorf 2004a; Miller etal. 2014;
Maharachchikumbura etal. 2015; Hyde etal. 2020). However,
Endophragmiella species were found to belong to Tubeufiales
and Helminthosphaeriaceae in several phylogenies (Hernán-
dez-Restrepo etal. 2017; Vu etal. 2019). Tengiomyces which
lacks DNA sequence data and phylogenies is more similar
to Chaetosphaerellaceae with ellipsoidal, septate ascospores
(Réblová 1999a; Huhndorf etal. 2004a).
Diaporthomycetidae was divided from Sordariomyceti-
dae based on DNA phylogenies and its members are wide-
spread as endophytic, pathogenic, parasitic or saprobic fungi
(Maharachchikumbura etal. 2015). Its members usually
have globose to subglobose ascomata, cylindrical to clavate
asci, variously shaped ascospores and hyaline, aseptate
conidia (Maharachchikumbura etal. 2015). Fifteen orders
with different characteristics are included in this subclass,
among which Calosphaeriales is similar to Togniniales in
having clavate asci borne from spicate ascogenous hyphae,
but are phylogenetically distant (Maharachchikumbura etal.
2016; Réblová etal. 2015; Hyde etal. 2020). The mono-
typic Togniniales was delimited based on molecular data
of Phaeoacremonium (Maharachchikumbura etal. 2016;
Hyde etal. 2020). Calosphaeriales comprises Calospha-
eriaceae and Pleurostomataceae, initially including eight
genera (Lumbsch and Huhndorf 2010; Maharachchikum-
bura etal. 2015, 2016). Later, Calosphaeria, Flabellascus,
Jattaea, Togniniella and Pleurostoma were confirmed to
belong to Calosphaeriales, but Sulcatistroma was nested in
Hypocreales (Vu etal. 2019; this study). The taxonomy of
Kacosphaeria and Tulipispora, which lack molecular data,
are rarely discussed.
Maharachchikumbura etal. (2015) established monotypic
Jobellisiales in Diaporthomycetidae, and then this order was
Fungal Diversity
1 3
listed in Hypocreomycetidae (Hyde etal. 2020; Wijaya-
wardene etal. 2020). However, Jobellisia species formed
an unstable monophyletic clade in Diaporthomycetidae
(Réblová 2008; Liu etal. 2012; Maharachchikumbura etal.
2015, 2016; Hongsanan etal. 2017; Hyde etal. 2020), and
they are similar to some Diaporthales species has yellow
to brown ascomata, cylindrical asci and brown ascospores
(Maharachchikumbura etal. 2015; Senanayake etal. 2017,
2018).
Morphology is the traditional method of delimiting spe-
cies. For the earlier and/or infrequently mentioned spe-
cies, they were initially classified mainly by morphology,
which has been demonstrated to be sometimes inconsistent
with DNA based phylogenies. García etal. (2004) and Cai
etal. (2006a, b) demarcated Neurospora species based on
the ascospores wall structure, but it was incongruent with
molecular data. Many species could not be accurately clas-
sified due to a lack of molecular or morphological informa-
tion. Therefore, it is necessary to determine phylogenetically
significant morphology coupled with DNA sequence data
to better infer species classification. Recently, Wang etal.
(2016, 2019a, b) and Marin-Felix etal. (2020) examined
several authentic materials and sequenced species in Chaeto-
miaceae and Lasiosphaeriaceae and delimited about twenty
genera and over fifty species based on phylogenetic analysis.
Vu etal. (2019) also sequenced a large number of species to
determine the taxonomic placement of more Sordariomy-
cetes species. We re-examined the type materials, authentic
specimens or extant literature pertaining to those species
that have been earlier/seldom mentioned or whose classifi-
cation is still controversial within the Diaporthomycetidae
and Sordariomycetidae. Wherever possible, DNA sequence
data are analyzed to update the taxonomy of these species.
Materials andmethods
Material examination
Specimens were loaned from BPI, BRIP, F, G, GZU, IMI,
NBRC, NY, PC, PDD, PRM, S, TRTC and UPS herbaria.1
These samples were observed in sterilized water and/or
5% KOH for sectioning. Specimens were examined using
a Motic SMZ 168 stereomicroscope. Micromorphologi-
cal 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 cam-
era. Tarosoft ® Image Framework program version 0.9.7
was used for measurements (Senanayake etal. 2020). Type
specimens those were in poor condition or unable to loaned,
morphological features based on the original publications
were re-drawn. The photoplates were processed with Adobe
Photoshop CS6 software (Adobe Systems, USA).
Phylogenetic analyses
Sequences were obtained from GenBank following previ-
ous publications and aligned using the default settings of
MAFFT version 7 (http:// mafft. cbrc. jp/ align ment/ server/
index. html) (Katoh and Standley 2013). These datasets were
manually corrected using Bioedit version 7.0.9.0 (Hall 1999;
Dissanayake etal. 2020). The phylogenetic analyses of the
combined gene regions were performed using maximum-
likelihood (ML) and/or Bayesian inference (BI). The evolu-
tionary model was obtained using jModeltest version 2.1.1
(Darriba etal. 2012) or ModelFinder (Kalyaanamoorthy
etal. 2017). The TIM1, TIM2, TIM3, TrN and TrNef sub-
stitution models were replaced by the GTR model (Lecocq
etal. 2013). The ML analyses were run with IQ-TREE
(Nguyen etal. 2015; Chernomor etal. 2016), using models
for each partitioned gene, with 1000 rapid bootstrap repli-
cates. BI was implemented by MrBayes v.3.0b4 (Ronquist
and Huelsenbeck 2003) with the best-fit model of sequence
evolution. Phylogenetic trees were viewed with FigTree
v1.4.0 (http:// tree. bio. ed. ac. uk/ softw are/ figtr ee/) and pro-
cessed by Adobe Illustrator CS5. The final alignments and
trees were deposited in TreeBASE (Tables2,3,4,5,6,7).
Diaporthomycetidae Senan., Maharachch. & K.D. Hyde,
Fungal Diversity 72: 208 (2015)
Maharachchikumbura etal. (2015) introduced the sub-
class Diaporthomycetidae, which is related to Sordariomy-
cetidae based on phylogenetic analyses (Hongsanan etal.
2017; Hyde etal. 2017, 2020; this study). There are 15
orders in this subclass, viz. Annulatascales, Atractosporales,
Calosphaeriales, Diaporthales, Distoseptisporales, Jobel-
lisiales, Magnaporthales, Myrmecridiales, Ophiostoma-
tales, Pararamichloridiales, Phomatosporales, Sporidesmi-
ales, Tirisporellales, Togniniales and Xenospadicoidales
(Maharachchikumbura etal. 2015; Hyde etal. 2020; this
study, Fig.1) and they are widely distributed as saprobes,
pathogens, or endophytes in aquatic or/and terrestrial habi-
tats (Hongsanan etal. 2017; Hyde etal. 2017, 2020). The
known asexual morphs in this subclass have been linked to
hyphomycetous or coelomycetous fungi generally forming
1 Institution: BPI: U.S. National Fungus Collections, USDA-ARS,
USA; BRIP: Department of Agriculture and Fisheries, Australia; F:
Field Museum of Natural History, USA; G: Conservatoire et Jardin
botaniques de la Ville de Genève, Switzerland; GZU: Karl-Franzens-
Universität Graz, Austria; IMI: CABI Bioscience UK Centre, UK;
NBRC: National Institute of Technology and Evaluation, Japan;
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.
Fungal Diversity
1 3
phialidic conidiogenesis and holoblastic conidiogenous
cells (Zhang and Blackwell 2001; Vijaykrishna etal. 2004;
Réblová etal. 2004, 2015; Najwa etal. 2012; Senanayake
etal. 2017; Wijayawardene etal. 2017).
Calosphaeriales M.E. Barr, Mycologia 75(1): 11 (1983)
The genera in Calosphaeriales are characterized by black
ascomata with an eccentric papilla or long neck, with con-
spicuously septate, tapered paraphyses, numerous, clavate
or subglobose asci borne from fascicled or spicate ascog-
enous hyphae, hyaline, allantoid to subcylindrical ascospores
and phialidic hyphomycetous asexual morphs (Barr 1985;
Réblová etal. 2004, 2015; Réblová and Mostert 2007). Calo-
sphaeriales and Togniniales are similar in morphology, but
are phylogenetically distant (Réblová etal. 2015; this study,
Fig.1). Ascogenous hypha, on which the asci form, are con-
spicuous character of members in Calosphaeriales and Tog-
niniales, and is an important feature to distinguish the genera
in these two orders (Barr 1985; Mostert etal. 2006; Damm
etal. 2008; Réblová etal. 2015). Two families are included
in this order, Calosphaeriaceae with four genera, and the
monotypic Pleurostomataceae (Wijayawardene etal. 2020;
this study). Jobellisiaceae was transferred to Calosphaeriales
based on multigene analysis (Hongsanan etal. 2017). How-
ever, Hongsanan etal. (2017) mentioned that Jobellisiaceae
is an unstable group and it is closely related to Calosphaeri-
ales, Diaporthales and Togniniales (Hyde etal. 2017; this
study, Fig.1). We place Jobellisiaceae under Jobellisiales
(see notes for Jobellisiales). Calosphaeriales is sister to Tog-
niniales in Diaporthomycetidae (Fig.1).
Calosphaeriaceae Munk, Dansk bot. Ark. 17(no. 1): 278
(1957)
Saprobic on dead wood or leaves in aquatic and/or ter-
restrial habitats, sometimes fungicolous. Sexual morph:
Ascostromata semi-immersed to erumpent through the
surface of the host. Ascomata perithecial, gregarious or
scattered or solitary, immersed, semi-immersed or super-
ficial, globose to subglobose, brown to black, coriaceous
to membranaceous, tuberculate or smooth or with hyphal
coating, ostiolate, with long or short necks. Peridium coria-
ceous or membranaceous, composed of hyaline to brown
cells of textura angularis to textura prismatica. Paraphyses
numerous, broad, cylindrical, septate, tapered. Ascogenous
hyphae discrete, hyaline, smooth-walled, branched, with
obovoid to obpyriform cells formed in sympodial succes-
sion, growing in dense clusters and each giving rise to an
ascus. Asci 8-spored, unitunicate, clavate to subglobose,
pedicellate, apex rounded or truncate, with or without an
apical ring. Ascospores bi-seriate to overlapping, allantoid
to ellipsoidal, straight to slightly curved, mostly aseptate,
sometimes multi-septate and producing numerous ascoco-
nidia when mature, hyaline to pale brown, smooth-walled,
sometimes with guttules. Ascoconidia hyaline, allantoid
to ellipsoidal, aseptate, smooth-walled, guttulate. Asexual
morph: Hyphomycetous. Conidiophores micronematous
or mononematous, hyaline, straight or flexuous, septate,
branched or unbranched. Phialides enteroblastic or holo-
blastic, monophialidic, hyaline, smooth-walled. Conidia
aggregated, hyaline, aseptate, cylindrical to allantoid, with
or without guttules (adapted from Réblová etal. 2004, 2015,
2016; Maharachchikumbura etal. 2016).
Type genus: Calosphaeria Tul. & C. Tul. 1863
Notes: Calosphaeria, Conidiotheca, Jattaea, Kacospha-
eria, Sulcatistroma, Togniniella and Tulipisopora have tradi-
tionally been accommodated Calosphaeriaceae (Maharach-
chikumbura etal. 2015, 2016; Wijayawardene etal. 2018).
Most species are wood inhabitants and are mostly reported
from Asia and Europe (Barr 1985; Damm etal. 2008;
Réblová etal. 2015). Réblová etal. (2004) established Calo-
sphaeriophora as a member of Calosphaeriaceae, typified
by Cal. pulchella. Subsequently, according to nomenclatu-
ral priority, Réblová etal. (2016) recommended that Calo-
sphaeriophora should be synonymized under Calosphaeria
(Réblová etal. 2004, 2015, 2016). Vu etal. (2019) sequenced
the ex-type strain of Sulcatistroma and found that it is more
closely related to Allantonectria (Nectriaceae) (Ramaley
2005; Hirooka etal. 2012; this study). Kacosphaeria was
accepted as a member of Calosphaeriaceae based on the sim-
ilarity of clavate asci with Calosphaeria (Spegazzini 1888).
We were unable to obtain authentic material of Kacospha-
eria and its description lacks sufficient detail to prove that
this genus belongs to Calosphaeriaceae. Therefore, we pro-
pose to transfer it to Calosphaeriales genera incertae sedis.
The Tulipispora has been placed in Calosphaeriaceae with-
out molecular data (Maharachchikumbura etal. 2015, 2016).
This hyphomycetous genus has multi-septate conidia which
is different from other members of Calosphaeriaceae. Gen-
era such as Triscelophorus and Triramulispora have multi-
septate conidia which are similar to Tulipispora (Révay etal.
2009). Thus, we place Tulipispora in Ascomycota genera
incertae sedis. The monotypic Conidiotheca is similar to
Jattaea and Pleonectria (Nectriaceae) in having ellipsoidal
to cylindrical ascospores producing ascoconidia (a conidium
formed directly from an ascospore, especially when still
within the ascus) (Hirooka etal. 2012). Molecular data of
Conidiotheca is unavailable and it has black ascomata and
muriform ascospores, instead of red ascomata in Pleonectria
or transverse-septate ascospores in Jattaea. Therefore, we
place this genus in Sordariomycetes genera incertae sedis
pending further studies. Réblová etal. (2015) introduced
Flabellascus as a member of Calosphaeriaceae based on
Fungal Diversity
1 3
Table 1 Change of taxa placement from Wijayawardene etal. 2020 to present
Taxa Taxa placement in Wijayawardene etal. 2020 Present placement
Jobellisiales M.J. D'souza & K.D. Hyde Hypocreomycetidae O.E. Erikss. & Winka Diaporthomycetidae Senan., Maharachch. &
K.D. Hyde
Apodospora Cain & J.H. Mirza Lasiosphaeriaceae Nannf Bombardiaceae S.K. Huang, Maharachch. &
K.D. Hyde
Apodus Malloch & Cain Lasiosphaeriaceae Nannf Neoschizotheciaceae S.K. Huang & K.D. Hyde
Arnium Nitschke ex G. Winter Lasiosphaeriaceae Nannf Sordariales genera incertae sedis
Biconiosporella Schaumann Lasiosphaeriaceae Nannf Sordariales genera incertae sedis
Bombardia (Fr.) P. Karst Lasiosphaeriaceae Nannf Bombardiaceae S.K. Huang, Maharachch. &
K.D. Hyde
Bombardioidea C. Moreau ex N. Lundqv Lasiosphaeriaceae Nannf Bombardiaceae S.K. Huang, Maharachch. &
K.D. Hyde
Boothiella Lodhi & Mirza Sordariomycetes genera incertae sedis Sordariaceae G. Winter
Camptosphaeria Fuckel Lasiosphaeriaceae Nannf Sordariales genera incertae sedis
Cercophora Fuckel Lasiosphaeriaceae Nannf Neoschizotheciaceae S.K. Huang & K.D. Hyde
Conidiotheca Réblová & L Mostert Togniniaceae Réblová, L. Mostert, W. Gams &
Crous Sordariomycetes genera incertae sedis
Copromyces N. Lundq Sordariaceae G. Winter Ascomycota genera incertae sedis
Diffractella Guarro etal Lasiosphaeriaceae Nannf Sordariales genera incertae sedis
Echria (N. Lundq.) Kruys etal Not mentioned Neoschizotheciaceae S.K. Huang & K.D. Hyde
Effetia Bartoli etal Sordariaceae G. Winter Ascomycota genera incertae sedis
Emblemospora Jeng & J.C. Krug Lasiosphaeriaceae Nannf Sordariales genera incertae sedis
Endophragmiella B. Sutton Helminthosphaeriaceae Samuels, Cand. &
Magni Ascomycota genera incertae sedis
Eosphaeria Höhn Lasiosphaeriaceae Nannf Sordariales genera incertae sedis
Episternus Górz & Boroń Lasiosphaeriaceae Nannf Zygospermellaceae S.K. Huang, Maharachch. &
K.D. Hyde
Fimetariella N. Lundq Lasiosphaeriaceae Nannf Bombardiaceae S.K. Huang, Maharachch. &
K.D. Hyde
Immersiella A.N. Mill. & Huhndorf Lasiosphaeriaceae Nannf Neoschizotheciaceae S.K. Huang & K.D. Hyde
Jugulospora N. Lundq Lasiosphaeriaceae Nannf Neoschizotheciaceae S.K. Huang & K.D. Hyde
Lasiosphaeris Clem Sordariales genera incertae sedis Lasiosphaeridaceae S.K. Huang, Maharachch. &
K.D. Hyde
Periamphispora J.C. Krug Lasiosphaeriaceae Nannf Sordariales genera incertae sedis
Ramophialophora M. Calduch etal Lasiosphaeriaceae Nannf Bombardiaceae S.K. Huang, Maharachch. &
K.D. Hyde
Rinaldiella Deanna A. Sutton etal Lasiosphaeriaceae Nannf Neoschizotheciaceae S.K. Huang & K.D. Hyde
Stellatospora T. Ito & Nakagiri Sordariaceae G. Winter Chaetomiaceae G. Winter
Strattonia Cif Lasiosphaeriaceae Nannf Strattoniaceae S.K. Huang, Maharachch. & K.D.
Hyde
Sulcatistroma A.W. Ramaley Calosphaeriales genera incertae sedis Hypocreales genera incertae sedis
Synaptospora Cain Helminthosphaeriaceae Samuels, Cand. &
Magni Sordariales genera incertae sedis
Tengiomyces Réblová Helminthosphaeriaceae Samuels, Cand. &
Magni Coronophorales genera incertae sedis
Tripterosporella Subram. & Lodha Lasiosphaeriaceae Nannf Sordariales genera incertae sedis
Tulipispora Révay & J. Gönczöl Sordariomycetes genera incertae sedis Ascomycota genera incertae sedis
Zygopleurage Boedijn Lasiosphaeriaceae Nannf Neoschizotheciaceae S.K. Huang & K.D. Hyde
Zygospermella Cain Lasiosphaeriaceae Nannf Zygospermellaceae S.K. Huang, Maharachch. &
K.D. Hyde
Fungal Diversity
1 3
morphological and phylogenetic analyses. Available phylo-
genetic data for Calosphaeriaceae, including Calosphaeria,
Flabellascus, Jattaea and Togniniella, are shown in Fig.2
and forms a sister clade to Pleurostomaceae in Calosphaeri-
ales (Fig.1).
Calosphaeria Tul. & C. Tul., Select. fung. carpol. (Paris)
2: 108 (1863)
Saprobic on dead wood. Sexual morph: Ascomata peri-
thecial, gregarious or solitary, immersed to semi-immersed,
globose to subglobose, brown to black, coriaceous to mem-
branaceous, tuberculate or smooth or with hyphal coating.
Necks long or short, oblong to subglobose, ostiolate, peri-
physate. Peridium comprising two layers, outer layer coria-
ceous, composed of brown cells of textura angularis; inner
layer membranaceous, composed of hyaline cells of textura
prismatica. Paraphyses numerous, broad, cylindrical, sep-
tate, longer than the asci. Ascogenous hyphae discrete, hya-
line, smooth-walled, branched, with ovoid to ellipsoidal
cells, growing in sympodial succession, in dense clusters
and each giving rise to an ascus. Asci 8-spored, unituni-
cate, clavate, pedicellate, apex rounded or truncate, with or
without J- apical ring. Ascospores bi-seriate to overlapping
in the upper part of ascus, allantoid to ellipsoidal, straight
to slightly curved, aseptate, hyaline, smooth-walled, some-
times with guttules. Asexual morph: Hyphomycetous. Con-
idiophores micronematous, hyaline, straight or flexuous,
septate, branched. Phialides enteroblastic or holoblastic,
monophialidic, elongate ampulliform to subcylindrical, hya-
line, smooth-walled. Conidia aggregated, hyaline, aseptate,
cylindrical to allantoid, with or without guttules (adapted
from Barr etal. 1993; Réblová etal. 2004).
Notes: Calosphaeria has been included in various fami-
lies, such as Diatrypaceae and Gnomoniaceae, based on the
tiny asci and allantoid ascospores (Barr etal. 1993; Réblová
etal. 2004). Tulasne and Tulasne (1863) established Calo-
sphaeria, typified by C. princeps. Over 80 species of Calo-
sphaeria have been found on wood in freshwater and/or
terrestrial habitats in America, Asia, Australia and Europe
(Barr 1985; Réblová etal. 2004; Damm etal. 2008), but few
have molecular data. Damm etal. (2008) and Réblová etal.
(2004) sequenced C. africana and C. pulchella, and these
Calosphaeria species form a sister clade to Jattaea in Calo-
sphaeriaceae based on multi-gene analysis (0.98PP, Fig.2).
Type species: Calosphaeria princeps Tul. & C. Tul.,
Select. fung. carpol. (Paris) 2: 109 (1863)
Basionym: Sphaeria pulchella Pers., Neues Mag. Bot.
1: 83 (1794)
Facesoffungi number: FoF 01135; Fig.3
Saprobic on dead wood. Sexual morph: Asco-
mata 300–450 µm (n = 10) diam. at the venter, peri-
thecial, gregarious, immersed, the long necks converg-
ing radially and erumpent through the bark of the host,
globose, brown to dark brown, coriaceous, glabrous. Necks
1.5–2.5 × 0.1–0.2mm (x̄ = 2 × 0.15mm, n = 10), coriaceous,
central, fasciculate, dark brown to black, smooth, wall com-
posed of brown to hyaline cells of textura epidermoidea to
textura prismatica (45–75µm wide, n = 20), ostiole with
hyaline periphyses. Peridium 40–80µm (x̄ = 55µm, n = 30)
wide at venter-walled, comprising two layers, outer layer
coriaceous, composed of brown cells of textura angularis;
inner layer membranaceous, composed of hyaline cells of
textura prismatica. Paraphyses 2–4.5µm (x̄ = 3.5µm, n = 30)
wide, numerous, cylindrical, septate, slightly constricted at
the septa, branched, apically rounded, tapering, longer than
the asci. Ascogenous hyphae discrete, elongate, hyaline,
smooth-walled, branched, with hyaline, ovoid to ellipsoi-
dal cells, 10–25 × 1.5–4µm (x̄ = 15.5 × 3µm, n = 20), grow-
ing in sympodial succession, in dense clusters, each giving
rise to an ascus. Asci 30–45(–50) × 3–5µm (x̄ = 40 × 4 µm,
n = 50), 8-spored, unitunicate, clavate, very long pedi-
cellate, apex blunt, without an apical ring. Ascospores
3.5–5.5 × 0.5–1.5µm (x̄ = 4 × 1µm, n = 50), overlapping
near the apex, hyaline, allantoid to ellipsoidal, straight to
slightly curved, aseptate, smooth-walled, sometimes with
guttules. Asexual morph: Hyphomycetous. Conidiophores
micronematous, hyaline, straight or flexuous, septate. Phi-
alides terminal or intercalary, monophialidic, ampulliform
to subcylindrical, hyaline, smooth-walled. Conidia hyaline,
aseptate, cylindrical to allantoid, with guttules (adapted from
Réblová etal. 2004).
Material examined: USA, Kansas, Stockton, on the inner
bark of Prunus cerasus, 5 October 1906, E. Bartholomew
(S-F263455); USA, Missouri, Emma, on dead trunks of Per-
sica vulgaris, November 1903, C.H. Demetrio (S-F263457);
France, 86 Vienne, Poitiers, Pictavii, on trunks of cherry
tree, 1840 (PC-MNHN-PC-PC0167696); France, 49 Maine-
et-Loire, Angers, on trunks of cherry tree, J.P. Guépin
(PC-MNHN-PC-PC0167711).
Known hosts and distribution: On decayed wood of Bet-
ula, Cerasus and Prunus in France (type locality), Germany
and North America (Persoon 1794; Fries 1823a; Tulasne and
Tulasne 1863; Schröter 1897; Barr 1985); on branches of
Solanum auriculatum in South Africa (Doidge 1950).
Notes: Persoon (1794) introduced Sphaeria pulchella
collected from decayed wood of Cerasus and Prunus and
Fries (1823a) made further collections on Betula and Prunus
in America. Tulasne and Tulasne (1863) established Calo-
sphaeria and transferred Sphaeria pulchella to Calosphaeria
princeps as the generic type. Schröter (1897) subsequently
re-named C. princeps as Calosphaeria pulchella as type
species, but this view was refuted by Clements and Shear
(1931) and Barr (1985). Damm etal. (2008) reported that
the ascospore size of ‘Calosphaeria princeps’ differs from
‘Calosphaeria pulchella’. In this study, based on priority, the
oldest name Calosphaeria princeps is recommended for use.
Fungal Diversity
1 3
Fig. 1 Phylogram generated
from maximum likelihood
analysis based on combined
LSU, SSU, RPB2 and TEF
sequence data of Sordariomy-
cetes. The confidence values
of bootstrap (BS) proportions
from the Maximum Likelihood
(ML) analysis (ML-BS > 50%)
above corresponding nodes. A
total of 139 strains are included
in the combined analyses, which
comprise 3892 characters (864
characters for LSU, 1049 char-
acters for SSU, 1059 characters
for RPB2, 920 characters for
TEF) after alignment. Strains of
Eurotiomycetes are used as out-
group taxa. The model of each
partitioned gene is GTR + I + G.
The best score in IQ-TREE
explores with a final likelihood
value of −65617.0325 is pre-
sented. Alignments are available
at TreeBASE (URL: http:// purl.
org/ phylo/ treeb ase/ phylo ws/
study/ TB2: S28266)
0.2
Coccodiella miconiae ppMP1342
Buergenerula spartinae ATCC 22848
Brunneodinemasporium jonesii GZCC_16-0050
Graphostroma platystoma AFTOL-ID 1249
Niesslia stellenboschiana CBS 145531
Phaeoacremonium novae-zealandiae WIN 113BI
Cladorrhinum foecundissimum CBS 180.66
Pararamichloridium caricicola CBS 145069
Podospora fimicola CBS 482.64
Monosporascus cannonballus MC1103
Triangularia bambusae CBS 352.33
Apiorhynchostoma curreyi UAMH 11088
Niesslia fuegiana CBS 368.77
Cryptosphaerella elliptica SMH4722
Lasiosphaeria sorbina CBS 885.85
Diatrype palmicola MFLUCC 110018
Falcocladium thailandicum CBS 121717
Plectosphaerella cucumerina CBS 131739
Lasiosphaeria ovina CBS 126299
Linocarpon cocois MFLU 15-2345
Neurospora endodonta IMI 148369
Ophiocordyceps sinensis YN09 64
Pseudovalsaria ferruginea UAMH 11490
Furfurella nigrescens CBS 143622
Sordaria prolifica CBS 567.72
Monosporascus nordestinus CMM-4846
Diatrypella japonica LL547
Biscogniauxia nummularia MUCL51395
Phialemonium obovatum CBS 279.76
Jobellisia guangdongensis HMAS 251240
Helminthosphaeria triseptata JF04015
Scortechiniellopsis leonensis GKM1269
Eutypa lata CBS 208.87
Microascus trigonosporus AFTOL-ID 914
Lanspora coronata AFTOL-ID 736
Pseudodactylaria brevis MFLUCC 16-0032
Xylaria hypoxylon CBS 122620
Meliola centellae VIC 31244
Falcocladium sphaeropedunculatum CBS 111292
Phomatospora biseriata MFLUCC 14-0832A
Plagiostoma aesculi AFTOL-ID 1238
Pseudoneurospora canariensis CBS 135818
Fragosphaeria reniformis CBS 134.34
Sulcatistroma nolinae CBS 117709
Juncigena adarca JK5548A
Robillarda sessilis CBS 114312
Myrmecridium iridis CPC 25084
Sporidesmium submersum MFLUCC 15-0421
Swampomyces armeniacus JK5325A
Ophiostoma piliferum AFTOL-ID 910
Pseudodactylaria camporesiana KUMCC 19-0074
Neospadicoides lignicola MFLUCC 17-2444
Ruzenia spermoides CBS 101621
Monosporascus ibericus CBS 110550
Furfurella stromatica CBS 144409
Linocarpon arengae MFLU 15-0306
Jobellisia fraterna SMH2863
Sporothrix brasiliensis CBS 133022
Spadicoides bina CBS 137794
Endothia gyrosa AFTOL-ID 1223
Chaetomium tenue CBS 139.38
Camarops tubulina SMH4614
Jattaea aurea CBS 140209
Coccodiella melastomatum CMU78543
Submersisphaeria aquatica A95-1B
Fragosphaeria purpurea CBS 133.34
Neofracchiaea callista SMH2689
Annulatascus velatisporus MFLU 16-2204
Calyptosphaeria tenebrosa PRA 12740
Calosphaeria africana STE-U 6182
Gliocephalotrichum bulbilium ATCC 22228
Ophiostoma saponiodorum CBS 128125
Sporidesmium fluminicola MFLUCC 15-0346
Pseudolachnella brevicoronata HHUF 30119
Allantonectria miltina CBS 125499
Monosporascus caatinguensis CMM-4833
Monosporascus mossoroensis CMM-4857
Conlarium aquaticum MFLUCC 15-0992
Malaysiasca phaii CBS 141321
Pseudodactylaria xanthorrhoeae CBS 143414
Ascocodinaea stereicola GJS95-184
Neopseudolachnella acutispora HHUF_29727
Phialemonium inflatum CBS 259.39
Coniochaeta luteoviridis CBS 206.38
Jobellisia luteola SMH2753
Phaeoacremonium fraxinopennsylvanica M.R. 3064
Phlogicylindrium uniforme CBS 131312
Ellisembia leonensis HKUCC 10822
Chaetomium globosum CBS 160.62
Monilochaetes infuscans CBS 869.96
Cordana inaequalis CBS 508.83
Endomeliola dingleyae PDD 98304
Xylaria multiplex HAST 580
Myrmecridium banksiae CPC 19852
Pseudoproboscispora caudae-suis A336-2D
Exophiala salmonis AFTOL-ID 671
Cryptosporella hypodermia AFTOL-ID 2124
Helminthosphaeria clavariarum ANM Acc.17
Delonicicola siamense MFLUCC 15-0670
Seiridum phylicae CPC 19962
Asteridiella obesa VIC 31239
Tirisporella beccariana BCC38300
Monosporascus semiaridus CMM-4830
Myrmecridium montsegurinum JF 13180
Monosporascus brasiliensis CMM-4839
Phomatospora bellaminuta AFTOL-ID 766
Dermatocarpon miniatum AFTOL-ID 91
Pleurostoma ochraceum CBS 131321
Doratomyces stemonitis AFTOL-ID 1380
Chaetomium pseudoglobosum CBS 574.71
Pleurostoma richardsiae CBS 270.33
Nakataea oryzae M21
Ceratocystiopsis minuta CBS 116963
Pararamichloridium verrucosa CBS 128.86
Falcocladium turbinatum BCC22055
Phialemonium atrogriseum CBS 604.67
Thailandiomyces bisetulosus BCC00018
Cordana pauciseptata IMI 232041a
Petriella setifera AFTOL-ID 956
Bussabanomyces longisporus CBS 125232
Etheirophora blepharospora JK5397A
Torrubiella wallacei CBS 101237
Ellisembia adscendens HKUCC 10820
Diatrype disciformis AFTOL-ID 927
Thyronectria concentrica ALLA
Leptosporella bambusae MFLUCC 12-0846
Nectria balansae AR4635
Sporothrix schenckii CBS 130114
Pleurostoma repens CBS 294.39
Rubellisphaeria abscondita CBS 132078
Lasiosphaeria lanuginosa SMH3819
Monosporascus eutypoides MT 45
Exophiala dermatitidis AFTOL-ID 668
Pseudoproboscispora thailandensis MFLUCC 15-0989
Phyllachora graminis UME 31349
Jattaea mookgoponga CBS 120867
Xylaria adscendens JDR 865
Exophiala pisciphila AFTOL-ID 669
Leptosporella arengae MFLUCC 15-0330
100
94
100
99
71
100
89
52
93
99
94
100
99
98
100
100
100
91
100
96
67
84
100
100
83
100
73
100
82
63
100
100
83
89
100
83
52
76
58
100
58
57
97
71
99
100
100
61
77
100
100
95
95
62
68
76
66
100
72
81
100
100
96
89
72
95
100
97
75
88
59
77
100
65
100
93
100
100
100
81
99
99
92
100
99
84
57
96
100
100
73
87
55
100
64
100
92
70
74
93
68
78
100
100
77
57
65
90
100
100
100
100
100
70
100
95
83
63
92
92
Xenospadicoidales
Atractosporales
Sporidesmiales
Annulatascales
Myrmecridiales
Ophiostomatales
Magnaporthales
Distoseptisporales
Phomatosporales
Pararamichloridiales
Jobellisiales
Tirisporellales
Diaporthales
Togniniales
Pleurostomaceae
Calosphaeriaceae
Calosphaeriales
Diaporthomycetidae
Sordariaceae
Lasiosphaeriaceae
Chaetomiaceae
Podosporaceae
Phyllachorales
Boliniales
Pseudodactylariales
Coniochaetales
Cephalothecales
Meliolales
Helminthosphaeriaceae
Linocarpaceae
Chaetosphaeriaceae
Leptosporellaceae
Diatrypaceae
Xylariaceae
Graphostromataceae
Amphisphaeriales
Delonicicolales
Falcocladiales
Coronophorales
Torpedosporales
Hypocreales
Microascales
Glomerellales
Eurotiomycetes
Sordariales
Chaetosphaeriales
Xylariomycetidae
Hypocreomycetidae
Xylariales
Sordariomycetidae
Fungal Diversity
1 3
The type material could not be obtained. Therefore, we re-
examined the authentic samples collected from France and
the USA. They are morphologically similar to C. princeps as
described by Tulasne and Tulasne (1863). Phylogenetically,
in this study, C. pulchella (strain CBS 115999) is considered
as C. princeps and is sister to C. africana (100%ML/1.00PP,
Fig.2). However, more fresh collections and phylogenetic
information are expected to determine the affinities of Calo-
sphaeria species.
Flabellascus Réblová, in Réblová, Jaklitsch, Réblová&
Štěpánek, PLoS ONE 10(12): e0144616, 15 (2015)
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, immersed to semi-immersed, subglobose, ostiolate,
with elongate necks, periphysate. Peridium coriaceous to
membranaceous, composed of brown to hyaline cells of
textura angularis to textura prismatica. Paraphyses numer-
ous, broad, cylindrical, septate, tapering, longer than the
asci. Ascogenous hyphae discrete, hyaline, smooth-walled,
elongate, branched, with ellipsoidal to obpyriform cells,
growing in sympodial succession, in dense clusters and
each giving rise to an ascus. Asci 8-spored, unitunicate,
clavate, pedicellate, thickened at the apex. Ascospores bi-
seriate to overlapping in the upper part of ascus, hyaline,
allantoid to ellipsoidal, smooth-walled, aseptate. Asexual
morph: Hyphomycetous. Conidiophores macronematous
to semi-macronematous, brown, unbranched or branched,
with terminate in a single or multiple verticillate phialide(s).
Phialides ampulliform to elongate-ampulliform, hyaline to
pale brown, tapering. Conidia aggregated, hyaline, aseptate,
allantoid to reniform (adapted from Réblová etal. 2015).
Notes: Flabellascus is monotypic. Its sexual morph is
similar to Jattaea and different asexual morphs distinguish
them. The former has macronematous and branched con-
idiophores, whereas the latter generally has micronematous
and unbranched conidiophores (Réblová etal. 2015). In our
study, Flabellascus is sister to Togniniella in Calospha-
eriaceae (100%ML/1.00PP, Fig.2).
Type species: Flabellascus tenuirostris Réblová, in
Réblová, Jaklitsch, Réblová & Štěpánek, PLoS ONE 10(12):
e0144616, 15 (2015)
Facesoffungi number: FoF 10116; Fig.4a–b
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, immersed, the necks converging radially and erumpent
0.1
Jattaea mucronata MFLUCC 16-0534
Phialemonium obovatum CBS:279.76
Jattaea mookgoponga CBS:120867
Pleurostoma richardsiae CBS 270.33
Jattaea discreta CBS 127681
Flabellascus tenuirostris CBS 138690
Flabellascus tenuirostris CBS 138692
Pleurostoma repens CBS 294.39
Phialemonium inflatum CBS:259.39
Jattaea aurea CBS 140209
Pleurostoma ochraceum CBS 131321
Calosphaeria africana CBS:120870
Phialemonium atrogriseum CBS:604.67
Jattaea prunicola CBS 120871
Jattaea tumidula CBS 140208
Jattaea taediosa PRM 934412
Jattaea aphanospora PRM 934328
Jattaea bruguierae MFLU:17-2648
Calosphaeria princeps CBS 115999
Pleurostoma ootheca CBS 115329
Togniniella microspora CBS 113726
Jattaea ribicola CBS 139779
Togniniella microspora CBS 125298
Jattaea leucospermi CBS:119343
Flabellascus tenuirostris CBS 138680
100/1.00
100/1.00
8 8/--
73/--
100/1.00
100/1.00
76/1.00
9 8/1.00
99/1.00
100/1.00
100/1.00
80/--
100/1.00
9 6/1.00
100/1.00
72/1.00
--/0.98
88/1.00
100/1.00
84/0.98
100/1.00
100/1.00
9 9/1.00
Calosphaeriaceae
Pleurostomaceae
Calosphaeriales
Cephalothecales
Fig. 2 Phylogram generated from maximum likelihood analysis based
on combined LSU, SSU, ITS, TUB and RPB2 sequence data of Calo-
sphaeriales. The confidence values of bootstrap (BS) proportions
from the Maximum Likelihood (ML) analysis (ML-BS > 70%, before
the backslash) and the posterior probabilities (PP) from the Bayesian
(BY) analysis (BY-PP > 0.95, after the backslash) above correspond-
ing nodes. The ‘--’ indicates lack of statistical support (< 70% for
ML-BS and < 0.95 for BY-PP). Twenty-five strains are included in the
combined analyses, which comprise 4592 characters (991 characters
for LSU, 1596 characters for SSU, 478 characters for ITS, 475 char-
acters for TUB, 1052 characters for RPB2) after alignment. Strains
of Coniochaetales are used as the outgroup taxa. The best score in
IQ-TREE explores with a final likelihood value of −21774.1640 is
presented. The model of each partitioned gene is: LSU: GTR + I + G;
SSU: GTR + I + G; ITS: GTR + I + G; TUB: HKY + I + G; RPB2:
GTR + I + G. The strain numbers are noted after the species names.
Ex-type strains are in bold. Alignments are available at TreeBASE
(URL: http:// purl. org/ phylo/ treeb ase/ phylo ws/ study/ TB2: S28265)
Fungal Diversity
1 3
through the bark of the host, subglobose, ostiolate, with
elongate necks, periphysate. Peridium coriaceous to mem-
branaceous, composed of brown to hyaline cells of textura
angularis to textura prismatica. Paraphyses numerous, broad,
cylindrical, septate, tapering, longer than the asci. Ascogenous
hyphae discrete, hyaline, smooth-walled, elongate, branched,
with ellipsoidal to obpyriform cells, growing in sympodial
succession, in dense clusters and each giving rise to an ascus.
Asci 8-spored, unitunicate, clavate, pedicellate, conspicuously
thickened at the apex. Ascospores bi-seriate to overlapping
in the upper part of ascus, hyaline, allantoid to ellipsoidal,
smooth-walled, aseptate. Asexual morph: Hyphomycetous.
Conidiophores macronematous to semi-macronematous,
brown, unbranched or branched, with terminate in a single
Fig. 3 Calosphaeria prin-
ceps: a, i, l, m (PC-MNHN-
PC-PC0167711); b–g, j,
k, p (S-F263455); h, q
(S-F263457); n, o (PC-MNHN-
PC-PC0167696); r (redrawn
from Réblová etal. 2004). a
Herbarium material. b, c Gre-
garious ascomata. d Ascoma.
e, f Ascoma in cross section.
g Wall of neck. h Numerous
asci and paraphyses. i Peridium
on the basal part of ascoma. j
Immature asci with branched
paraphyses. k Immature asci,
and paraphyses attached to
ascogenous hyphae with ovoid
to ellipsoidal cells (arrow
indicates a cell on ascogenous
hyphae). l–p Mature asci (n–p
stained in Melzer’s reagent).
q Ascospores. r Hyphae with
phialides and conidia. Scale
bars: b–c = 1mm, d–f = 200µm,
h. i = 100µm, j = 50µm, g.
k = 20µm, l–p. r = 10µm,
q = 2µm
Fungal Diversity
1 3
phialide or 2–3 verticillate phialides. Phialides ampulliform
to elongate-ampulliform, hyaline to pale brown, gradually
tapering. Conidia aggregated, hyaline, aseptate, allantoid to
reniform (adapted from Réblová etal. 2015).
Known hosts and distribution: On wood of Fagus syl-
vatica and Quercus cerris in Czech Republic (type locality)
(Réblová etal. 2015).
Notes: Réblová etal. (2015) introduced the characteris-
tics of Flabellascus tenuirostris in detail, and analyzed four
strains with ITS, LSU, TUB and RPB2 regions. The draw-
ing is provided for the type species (Fig.4 a–b), based on
Réblová etal. (2015).
Jattaea Berl., Icon. fung. (Abellini) 3(1–2): 6 (1900)
Saprobic on wood. Sexual morph: Ascomata perithecial,
aggregated to scattered, immersed to semi-immersed, often
associated with other dead ascomata, brown to dark brown,
globose to subglobose, membranaceous, glabrous or covered
with hyphae, ostiolate, with long or short necks, periphysate.
Peridium comprising two layers, outer layer composed of
brown to pale brown cells of textura angularis to textura
prismatica; inner layer composed of hyaline cells of textura
prismatica. Paraphyses numerous, broad, cylindrical, sep-
tate. Ascogenous hyphae discrete, hyaline, smooth-walled,
with ellipsoidal to obpyriform cells, growing in sympo-
dial succession, in dense clusters and each giving rise to
an ascus. Asci 8-spored, unitunicate, clavate, pedicellate,
apex obtuse. Ascospores bi-seriate to overlapping, hyaline,
allantoid to ellipsoidal, aseptate to multi-septate, smooth-
walled, sometimes with guttules, sometimes producing
numerous ascoconidia when mature. Ascoconidia hyaline,
allantoid to ellipsoidal, aseptate, smooth-walled, guttulate.
Asexual morph: Hyphomycetous, Conidiophores hyaline
to pale brown, septate, unbranched to branched. Phialides
monophialidic, elongate to ampulliform, fusiform, smooth-
walled. Conidia hyaline, ovoid to suballantoid, straight or
slightly curved, aseptate, smooth-walled (adapted from Ber-
lese 1900; Réblová 2011).
Notes: Jattaea and Wegelina were proposed as inde-
pendent genera (Clements and Shear 1931). Subsequently,
Réblová (2011) showed that the type Wegelina discreta and
Jattaea species are congeneric based on DNA sequence
analyses, and reduced Wegelina to a synonym of Jattaea
(Réblová 2011). Jattaea resembles Calosphaeria, Flabel-
lascus, Pleurostoma and Togniniella in having clavate asci
and allantoid ascospores (Réblová 2011; Huang etal. 2018).
Jattaea species are aggregated and sister to Calosphaeria
(0.98PP) and distinct from Flabellascus, Pleurostoma and
Togniniella (Fig.2).
Type species: Jattaea algeriensis Berl., Icon. fung.
(Abellini) 3(1–2): 6 (1900)
Fig. 4 Flabellascus tenui-
rostris: a, b (redrawn from
Réblová etal. 2015). a Asci link
with ascogenous hyphae and
paraphyses. b Conidiophores
with conidia; Kacosphaeria
antarctica: c–e (redrawn from
Arambarri etal. 2020). c Asci
with paraphyses. d Ascospores.
e Ascomata. Scale bars:
a–d = 10µm, e = 200µm
Fungal Diversity
1 3
Facesoffungi number: FoF 10117
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, aggregated to scattered, immersed, black, glabrous,
ostiolate, with necks. Asci 8-spored, unitunicate, cylindri-
cal. Ascospores hyaline, allantoid, aseptate, smooth-walled.
Asexual morph: Hyphomycetous, phialophora-like.
Conidia hyaline, ovoid to suballantoid, aseptate, smooth-
walled (adapted from Berlese 1900; Damm etal. 2008).
Known hosts and distribution: On decayed stems or wood
of Rubus in Algeria (type locality) (Berlese 1900).
Notes: Jattaea algeriensis was reported on the sarmen-
tum of Rubus fruticosus in Algeria (Berlese 1900). Réblová
(2011) used CBS 120871 to stand for J. algeriensis. How-
ever, strain CBS 120871 was sequenced from type material
of Jattaea prunicola (Damm etal. 2008). The asci of J.
algeriensis is clavate with long pedicel which is different
from J. prunicola with cylindrical asci (Damm etal. 2008).
We were unable to obtain authentic materials or find a reli-
able illustration. Therefore, we re-examined lectotype of Jat-
taea discreta, which is similar to J. algeriensis (Damm etal.
2008; Réblová 2011), and provide an illustration. Fresh col-
lections and phylogenetic data of J. algeriensis are needed
to determine its taxonomic placement.
Jattaea discreta (Berl.) Réblová, Fungal Diversity 49: 179
(2011)
Basionym: Wegelina discreta Berl., Icon. fung. (Abellini)
3(1–2): 8 (1900)
Facesoffungi number: FoF 09998; Fig.5
Saprobic on wood. Sexual morph: Ascomata 200–350µm
(n = 10) diam. at the venter, perithecial, aggregated to scat-
tered, immersed, the elongate necks converging radially
and erumpent through the bark of the host, often associated
with other dead ascomata, brown to dark brown, globose
to subglobose, glabrous. Necks 100–150 × 400–700 μm
(x̄ = 130 × 550µm, n = 10), central, ostiolate, periphysate,
straight or slightly flexuous, rounded to slightly tapering at
the apex, the wall composed of dark brown to brown cells
of textura epidermoidea to textura prismatica. Peridium
20–55µm (x̄ = 35µm, n = 30) wide at venter-walled, com-
prising two layers, outer layer composed of brown to pale
brown cells of textura angularis to textura prismatica;
inner layer composed of hyaline cells of textura prismatica.
Paraphyses 1–2µm wide, numerous, broad, cylindrical,
septate, slightly constricted at the septa, unbranched, apex
rounded. Ascogenous hyphae discrete, hyaline, smooth-
walled, branched, with hyaline, ellipsoidal to obpyriform
cells, 3–4.5 × 1.5–2.5µm (x̄ = 3.5 × 2µm, n = 20), growing in
sympodial succession, in dense clusters and each giving rise
to an ascus. Asci (15–)17–23(–30) × 2–4µm (x̄ = 20 × 3µm,
n = 50), 8-spored, unitunicate, clavate, apex obtuse, with-
out apical ring, long pedicellate. Ascospores 4–6 × 1–2µm
(x̄ = 5 × 1.5µm, n = 50), bi-seriate to overlapping, hyaline,
allantoid to ellipsoidal, slightly curved, aseptate, smooth-
walled, sometimes with guttules. Asexual morph: Hypho-
mycetous. Conidiophores hyaline to pale brown, erect,
septate, unbranched or branched. Phialides monophialidic,
terminal to intercalary, elongate to ampulliform, fusiform,
narrowed at the apex, smooth-walled, hyaline, with cylindri-
cal to ampulliform adelophialides. Conidia hyaline, ovoid to
suballantoid, straight or slightly curved, aseptate, smooth-
walled, aggregated, accumulating in slimy masses on the
apex of the phialides (adapted from Réblová 2011).
Material examined: Italy, Veneto, Belluno, on decaying
wood of a branch of Acer pseudoplatanus, Autumn 1879,
(NY-00912077, lectotype).
Known hosts and distribution: On decayed wood of Cra-
taegus, Acer, Quercus, Vaccinium in France, Italy (type
locality) and USA (Berlese 1900; Réblová 2006, 2007,
2011).
Notes: Wegelina species have been transferred to Barbat-
osphaeria, Ceratostomella and Jattaea (Kummer etal. 2005;
Réblová 2006, 2007, 2011). Jattaea and Wegelina have
similar characters with clavate asci and hyaline, allantoid
ascospores, but differ in the length of the ascomata’s neck
(Berlese 1900). Réblová (2011) confirmed that Wegelina dis-
creta is a member of Jattaea. This sample (NY-00912077)
was initially determined as Calosphaeria wahlenbergii, then
identified as an isotype of Wegelina discreta by Barr in 1984
(mentioned in the label of material), and designated as a
lectotype of J. discreta by Réblová (2011).
Togniniella Réblová, L. Mostert, W. Gams & Crous, Stud.
Mycol. 50(2): 543 (2004)
Saprobic on wood. Sexual morph: Ascomata perithecial,
gregarious or solitary, immersed to semi-immersed, globose
to subglobose, glabrous, brown to black, carbonaceous to
coriaceous, immersed, the long necks erumpent through
the bark of the host. Necks central, cylindrical or slightly
flexuous, black, smooth, apex broadly rounded, ostiolate,
periphysate. Peridium comprising two layers, outer layer
carbonaceous or coriaceous, composed of brown cells of
textura angularis; inner layer membranaceous, composed of
hyaline cells of textura prismatica. Paraphyses unbranched,
broad, cylindrical, septate, slightly constricted at the septa,
tapering, with a rounded apex, longer than asci. Ascoge-
nous hyphae discrete, hyaline, smooth-walled, branched,
with hyaline, obovoid to ellipsoidal cells, growing in sym-
podial succession, in dense clusters, each giving rise to
an ascus. Asci 8-spored, unitunicate, clavate, pedicellate,
apex obtuse, without apical ring. Ascospores overlapping
near the apex, allantoid to suballantoid, straight to slightly
curved, aseptate, hyaline, smooth-walled, sometimes with
guttules. Asexual morph: Hyphomycetous. Conidiophores
macronematous or micronematous, usually arising from
aerial hyphae, erect, hyaline to pale brown, unbranched or
Fungal Diversity
1 3
branched, straight or flexuous, septate, constricted at the
septa. Phialides monophialidic or polyphialidic, terminal
or intercalary, subcylindrical to ampulliform, tapering,
smooth-walled, hyaline, with cylindrical adelophialides.
Conidia hyaline, obovoid to reniform, straight or slightly
curved, aseptate, smooth-walled, accumulating in slimy
masses on the apex of the phialides (adapted from Réblová
etal. 2004).
Notes: The monotypic Togniniella initially typified by T.
acerosa, and its asexual morph was introduced as Phaeo-
crella acerosa (Réblová etal. 2004). Subsequently, the name
T. acerosa was corrected as T. microspora (Réblová 2011;
Réblová etal. 2015). Réblová etal. (2016) recommended
that Phaeocrella should be synonymized under Togniniella,
according to nomenclatural priority. This genus is character-
ized by clavate, tiny asci, allantoid ascospores and verrucous
Fig. 5 Jattaea discreta: a–q (NY-00912077, lectotype); r (redrawn
from Réblová 2011). a Herbarium material. b–d Ascomata erumpent
through the bark of the host. e Ascoma in cross section. f Perid-
ium. g Transverse section of neck. h, i Asci linked with ascogenous
hyphae and paraphyses. j–m Asci. n–q Ascospores, r. Conidiophores
with conidia. Scale bars: b. c = 500 µm, d = 1 mm, e = 200 µm, f.
g = 50µm, h. i = 20µm, j–m, r = 10µm, n–q = 2µm
Fungal Diversity
1 3
phialides with reniform conidia (Réblová etal. 2004, 2015,
2016; Réblová 2011), and phylogenetic information is
available (Réblová etal. 2004; Damm etal. 2008; Vu etal.
2019). In this study, Togniniella is sister to Flabellascus
(100%ML/1.00PP, Fig.2).
Type species: Togniniella microspora (Ellis & Everh.)
Réblová, Fungal Diversity 49: 193 (2011)
Basionym: Ceratostomella microspora Ellis & Everh.,
Proc. Acad. nat. Sci. Philad. 45: 444 (1894)
Facesoffungi number: FoF 09999; Fig.6
Fig. 6 Togniniella microspora:
a, b, d–e, h–j (PDD-81431,
holotype); c, f, g (PDD-81432);
k, l (redrawn from Réblová
2011). a Herbarium material.
b, c Ascomata. d Peridium.
e Septate paraphysis. f–j
Asci. k Asci with ascogenous
hyphae. l. Conidiophores with
conidia. Scale bars: b = 200µm,
c = 500µm, d = 20µm, e. k.
l = 10µm, f–j = 5µm
Fungal Diversity
1 3
Saprobic on wood. Sexual morph: Ascomata
200–400µm (250µm, n = 10) diam. at the venter, perithe-
cial, gregarious or solitary, immersed to semi-immersed, the
long necks erumpent through the bark of the host, globose
to subglobose, glabrous, brown to black, carbonaceous to
coriaceous. Necks 550–1000 × 55–75µm (x̄ = 755 × 65µm,
n = 10), carbonaceous, central, cylindrical or slightly flex-
uous, black, smooth, apex broadly rounded, ostiole with
hyaline periphyses. Peridium 20–30µm (x̄ = 25µm, n = 30)
wide at venter-walled, comprising two layers, outer layer
carbonaceous to coriaceous, composed of brown cells
of textura angularis; inner layer membranaceous, com-
posed of hyaline cells of textura prismatica. Paraphyses
7–8µm (x̄ = 7.5µm, n = 30) wide near the base, 1.5–5µm
(x̄ = 2.5µm, n = 30) wide near the apex, unbranched, broad,
cylindrical, septate, slightly constricted at the septa, taper-
ing, with a rounded apex, longer than asci. Ascogenous
hyphae discrete, hyaline, smooth-walled, branched, with
hyaline, obovoid to ellipsoidal cells, 3–4.5 × 1.5–2.5µm
(x̄ = 3.5 × 2 µm, n = 20), growing in sympodial succes-
sion, in dense clusters, each giving rise to an ascus. Asci
10–16(–20) × 2–5µm (x̄ = 15 × 3µm, n = 50), 8-spored, uni-
tunicate, clavate, long pedicellate, apex obtuse, apical ring
absent. Ascospores 2.5–4 × 0.2–0.5µm (x̄ = 3.5 × 0.3 µm,
n = 50), overlapping near the apex, allantoid to suballantoid,
straight to slightly curved, aseptate, hyaline, smooth-walled,
sometimes with guttules. Asexual morph: Hyphomycetous.
Conidiophores macronematous or micronematous, usually
arising from aerial hyphae, erect, hyaline to pale brown,
unbranched or branched, straight or flexuous, 1–4-sep-
tate, constricted at the septa. Phialides monophialidic or
polyphialidic, terminal to intercalary, subcylindrical to
ampulliform, tapering, smooth-walled, hyaline, with cylin-
drical adelophialides. Conidia hyaline, obovoid to reniform,
straight or slightly curved, aseptate, smooth-walled, aggre-
gated into a round at the apex of phialides (adapted from
Réblová etal. 2004).
Material examined: New Zealand, South Island, Tasman
Province, Nelson Lake National Park, St. Arnaud, Lake
Rotoiti, Lakehead track ca. 1.5km SE of NP Headquarters,
on decayed wood of a trunk of Nothofagus sp., 22 Febru-
ary 2003, M. Réblová (PDD-81431, holotype); New Zea-
land, West Coast Province, Harihari 73km SW of Hokitika,
Saltwater Forest, Poerua River Valley, on decayed wood, 12
March 2003, M. Réblová (PDD-81432).
Known hosts and distribution: On decayed wood of
Nothofagus, Populus in Canada (type locality) and New
Zealand (Réblová etal. 2004; Réblová 2011).
Notes: Togniniella microspora is sister to Flabellascus
tenuirostris (Réblová etal. 2004; Damm etal. 2008; Vu
etal. 2019; this study, 100%ML/1.00PP, Fig.2). Ascomata
of 81,431 and 81,432 are fragile, almost no ascoma with a
long intact neck is seen, and most features are evanescent.
We provided photo of observable characters and complete
the description through Réblová etal. (2004).
Pleurostomataceae Réblová, L. Mostert, W. Gams & Crous,
Stud. Mycol. 50(2): 540 (2004)
Saprobic on wood and/or isolated from soil; parasitic in
humans. Sexual morph: Ascomata perithecial, gregarious
or solitary, immersed or superficial, globose to subglobose,
dark brown to black, glabrous, ostiolate, with papilla, peri-
physate. Peridium outer layer coriaceous, composed of dark
brown to pale brown cells of textura intricata to textura epi-
dermoidea to textura angularis; inner layer composed of
hyaline cells of textura prismatica. Paraphyses not observed.
Ascogenous hyphae discrete, hyaline, smooth-walled,
branched, with hyaline, ellipsoidal cells, growing in sym-
podial succession, in dense clusters, each giving rise to an
ascus. Asci multi-spored, unitunicate, reniform to ellipsoidal,
with blunt or rounded, thickened apex. Ascospores overlap-
ping, cylindrical to allantoid, straight to curved, aseptate,
hyaline, smooth-walled, sometimes with guttules. Asexual
morph: Hyphomycetous. Mycelium composed of branched,
septate, tuberculate or smooth-walled, hyaline to brown
hyphae. Conidiophores usually arising from aerial or sub-
merged hyphae, erect, septate, straight or flexuous, hyaline
to brown. Phialides monophialidic or polyphialidic, with
inconspicuous or flared collarettes, smooth-walled, hyaline,
cylindrical. Conidia hyaline, ovoid to suballantoid, straight
or slightly curved, aseptate, smooth-walled, accumulating
in slimy masses on the apex of the phialides (adapted from
Vijaykrishna etal. 2004).
Type genus: Pleurostoma Tul. & C. Tul. 1863
Notes: Tulasne and Tulasne (1863) established Pleu-
rostoma candollei as the generic type. Vijaykrishna etal.
(2004) introduced a second species P. ootheca and its asex-
ual morph Pleurostomophora ootheca. Subsequently, three
more species, Pleurostomophora ochraceum (human patho-
gen), Ple. repens and Ple. richardsiae, were confirmed with
molecular data (Vijaykrishna etal. 2004; Najwa etal. 2012).
Pleurostomataceae was proposed as an independent family
in Calosphaeriales with two genera Pleurostoma and Pleuro-
stomophora based on morphological and molecular analysis
(Réblová etal. 2004). Réblová etal. (2015, 2016) recom-
mended that the asexual Pleurostomophora be reduced as a
synonym to Pleurostoma based on phylogenetic analysis and
to maintain nomenclatural stability. The monotypic nature of
Pleurostomataceae is characterized by branched ascogenous
hyphae, allantoid ascospores and ovoid conidia, which is
similar to Calosphaeriaceae, but the former has short-necked
ascomata and reniform asci, and the latter has long-necked
ascomata and clavate asci. Pleurostomataceae was revised
based on morphology and phylogenetic analysis and it is
confirmed as a sister family to Calosphaeriaceae in Calo-
sphaeriales (100%ML/1.00PP, Fig.2).
Fungal Diversity
1 3
Pleurostoma Tul. & C. Tul., Select. fung. carpol. (Paris) 2:
247 (1863)
Saprobic on wood and/or isolated from soil; parasitic in
humans. Sexual morph: Ascomata perithecial, gregarious
or solitary, at first immersed, then becoming superficial,
globose to subglobose, dark brown to black, coriaceous,
glabrous, ostiolate, with papilla, periphysate. Peridium
comprising three layers, outer layer coriaceous, composed
of dark brown to pale brown cells of textura intricata to
textura epidermoidea; middle layer coriaceous, composed
of brown cells of textura epidermoidea to textura angularis;
inner layer membranaceous, composed of hyaline cells of
textura prismatica. Paraphyses not observed. Ascogenous
hyphae discrete, hyaline, smooth-walled, branched, with
hyaline, ellipsoidal cells, growing in sympodial succes-
sion, in dense clusters, each giving rise to an ascus. Asci
multi-spored, unitunicate, reniform to ellipsoidal, producing
bristle-like pedicel when separating the ascogenous hyphae,
with blunt or rounded and thickened apex. Ascospores over-
lapping, cylindrical to allantoid, straight to curved, aseptate,
hyaline, smooth-walled, sometimes with guttules. Asexual
morph: Hyphomycetous. Mycelium composed of branched,
septate, tuberculate or smooth-walled, hyaline to brown
hyphae. Conidiophores usually arising from aerial or sub-
merged hyphae, erect, septate, straight or flexuous, hyaline
to brown. Phialides monophialidic or polyphialidic, with
inconspicuous or flared collarettes, smooth-walled, hyaline,
cylindrical. Conidia hyaline, ovoid to suballantoid, straight
or slightly curved, aseptate, smooth-walled, accumulating
in slimy masses on the apex of the phialides (adapted from
Tulasne and Tulasne 1863; Vijaykrishna etal. 2004).
Notes: Pleurostoma is characterized by glabrous, black
ascomata, reniform to ellipsoidal asci with a thickened
apex, cylindrical to allantoid ascospores and a phialophora-
like asexual morph (Réblová etal. 2004, 2015, 2016;
Vijaykrishna etal. 2004; Najwa etal. 2012). Two asexual
species, Pleurostoma minimum and P. vibratile, were trans-
ferred to Phaeoacremonium based on phylogenetic analysis
(Gramaje etal. 2015). Thus, Pleurostoma accommodates
five species, three have been reported as asexual (P. repens,
P. richardsiae and P. ochraceum), one sexual (P. candol-
lei) and one holomorphic species (P. ootheca) (Tulasne and
Tulasne 1863; Vijaykrishna etal. 2004; Najwa etal. 2012;
Réblová etal. 2015).
Type species: Pleurostoma candollei Tul. & C. Tul. [as 'can-
dollii'], Select. fung. carpol. (Paris) 2: 247 (1863)
Facesoffungi number: FoF 01138; Fig.7
Saprobic on decorticated wood. Sexual morph: Asco-
mata 470–520 × 370–420 µm (x̄ = 500 × 400µm, n = 10),
perithecial, gregarious or solitary, semi-immersed or superfi-
cial, globose to subglobose, dark brown to black, coriaceous,
glabrous, ostiolate papilla, central, periphysate. Peridium
35–95µm (x̄ = 60µm, n = 30) wide, comprising three lay-
ers, outer layer coriaceous, composed of dark brown to pale
brown cells of textura intricata to textura epidermoidea;
middle layer coriaceous, composed of brown cells of textura
epidermoidea to textura angularis; inner layer membrana-
ceous, composed of hyaline cells of textura prismatica. Par-
aphyses not observed. Ascogenous hyphae discrete, hyaline,
smooth-walled, branched, with hyaline, ellipsoidal cells,
1.5–4.5 × 1.5–3µm (x̄ = 2.5 × 2µm, n = 20), growing in sym-
podial succession, in dense clusters, each giving rise to an
ascus. Asci (15–)18–25(–35) × 6.5–11.5µm (x̄ = 23 × 8.5µm,
n = 50), multi-spored, unitunicate, reniform to ellipsoidal,
producing bristle-like pedicel when separating the ascog-
enous hyphae, with rounded and asymmetrically thickened
apex. Ascospores 2.5–4 × 0.5–1.5µm (x̄ = 3 × 1µm, n = 50),
overlapping, cylindrical to allantoid, straight to curved,
aseptate, hyaline, smooth-walled, sometimes with guttules.
Asexual morph: Undetermined.
Material examined: France, 92 Hauts-de-Seine, Chaville,
on dead wood of Quercus sp., 8 March 1859 (PC-MNHN-
PC-PC0167640, holotype); USA, Massachusetts, Medford,
on inner surface of bark of standing beech tree, 20 April
1936, G.D. Darker (S-F21567).
Known hosts and distribution: On decayed and decor-
ticated wood of Quercus in France (type locality), North
America (Tulasne and Tulasne 1863; Bates etal. 2018).
Notes: Tulasne and Tulasne (1863) examined specimens
of rotting bark-less oak collected in Chaville, France in
the winter and introduced Pleurostoma candollei. We re-
examined the specimen, PC0167640, which was collected
on decorticated wood of Quercus at Chaville in March 1859.
This sample was named Sphaeria latericollis DC. and Spha-
eria spermoides, and finally determined as Pleurostoma can-
dollei (mentioned in the label of material). This information
is similar to the history of type material of P. candollei men-
tioned in Tulasne and Tulasne (1863). Thus, we re-examined
the authentic material of PC0167640 and designated that it
is the holotype of P. candollei.
Tulasne and Tulasne (1863) also observed a kind of vil-
liform, brown to greyish mycelium with branched conidi-
ophores and aggregated, terminal, slender, straight or curved
conidia in nature on the same specimen (Fig.7r). However,
cultural characteristics and phylogenetic analysis have not
confirmed the relationship between this hyphomycetous fun-
gus and P. candollei. We could not find any asexual taxon
on the materials. The hand-drawing provided for this hypho-
mycete (Fig.7r) is based on Tulasne and Tulasne (1863).
Calosphaeriales genera incertae sedis
Kacosphaeria Speg., Boln Acad. nac. Cienc. Córdoba
11(2): 214 (1887) [1888]
Saprobic on wood. Sexual morph Ascomata perithecial,
aggregated, discoid, immersed to erumpent through the
Fungal Diversity
1 3
Fig. 7 Pleurostoma candol-
lei: a, b, e–j, n (PC-MNHN-
PC-PC0167640, holotype);
c, d, k–m, o–q (S-F21567);
r (redrawn from Tulasne and
Tulasne 1863). a Herbarium
material. b–d Ascomata (d
Ascomata in cross section on
host). e, f Ascoma in cross
section. g–i Peridium (h outer
layer, i inner layer). j Asci arise
from single cells on ascogenous
hyphae (arrows indicate cells)
k Mature and immature asci.
l, m Asci with apical thicken-
ing and short pedicel (l arrow
indicates an asymmetrically
thickened apex, k arrow indi-
cates a bristle-like pedicel).
n–q Ascospores. r Villiform
hyphae and slender conidia on
host. Scale bars: b = 500µm,
c–f = 200µm, r = 50µm, g,
i, j = 20µm, h, k = 10µm, l,
m = 5µm, n = 2µm, o–q = 1µm
Fungal Diversity
1 3
bark of the host, black, glabrous, thick-walled, with short
necks. Asci 8-spored, unitunicate, clavate, long pedicellate,
apex obtuse, with long paraphyses. Ascospores bi-seriate,
allantoid, 1-septate, hyaline. Asexual morph Undetermined
(adapted from Spegazzini 1888).
Notes: The monotypic Kacosphaeria with clavate asci
and allantoid ascospores that is similar to some species of
Calosphaeria (Spegazzini 1888). The specific ascogenous
hyphae of Calosphaeriales is not mentioned in the descrip-
tion (Spegazzini 1888), nor is it found in the drawings of
Spegazzini (Arambarri etal. 2020). Thus, we propose that
Kacosphaeria be placed in Calosphaeriales genera incertae
sedis, and await more fresh samples pending.
Type species: Kacosphaeria antarctica Speg., Boln Acad.
nac. Cienc. Córdoba 11(2): 214 (1887) [1888]
Facesoffungi number: FoF 10118; Fig.4c–e
Saprobic on Ribisma gellanicae. Sexual morph Asco-
mata 280–320µm, perithecial, 4–9 ascomata aggregated,
discoid, immersed to erumpent through the bark of the host,
black, glabrous, with short necks. Asci 40–50 × 4–5 µm,
8-spored, unitunicate, clavate, long pedicellate, obtuse apex,
with long paraphyses. Ascospores 9–11 × 1.5µm, bi-seriate,
allantoid, 1-septate, hyaline. Asexual morph Undetermined
(adapted from Spegazzini 1888).
Known hosts and distribution: On decayed wood of Ribisma
gellanicae in Argentina (type locality) (Spegazzini 1888).
Notes: Kacosphaeria antarctica was found on dead
branches of Ribisma gellanicae in Ushuaia, Argentina, in
June 1882 (Spegazzini 1888). We were unable to obtain
authentic material of K. antarctica, but provide a drawing
(Fig.4c–e) based on Spegazzini's drawings from Arambarri
etal. (2020).
Jobellisiales M.J. D'souza & K.D. Hyde 2015
Réblová (2008) and Liu etal. (2012) showed that Jobelli-
sia species were closely related to members of Calosphaeri-
ales, Diaporthales and Togniniaceae based on LSU sequence
analysis. Maharachchikumbura etal. (2015) established the
monotypic order Jobellisiales in Diaporthomycetidae based
on the multi-gene analysis. The phylogenetic position of this
order is unstable and was previously thought to be related
to Calosphaeriales and Togniniales (Hongsanan etal. 2017;
Hyde etal. 2017). We find that Jobellisiales is basal to
Calosphaeriales, Togniniales, Diaporthales and Tirisporel-
lales (Fig.1). Jobellisiales is different from Calosphaeri-
ales and Togniniales with clavate asci and hyaline allantoid
ascospores, and different from Tirisporellales with cylindri-
cal asci and fusoid, falcate to lunate ascospores (Hyde etal.
2020). It is similar to Diaporthales, in having cylindrical to
ellipsoidal asci and brown, subglobose ascospores; however,
they are distantly related in phylogenetic analyses (Hyde
etal. 2020; this study, Fig.1).
Jobellisiaceae Réblová, Mycologia 100(6): 899 (2008)
Saprobic on wood. Sexual morph: Ascostromata crus-
tose. Ascomata perithecial, gregarious, semi-immersed to
superficial, globose to subglobose, black to dark brown,
rough, ostiolate, with papilla, periphysate. Peridium mem-
branaceous, composed of blue-green or brown or yellow
to hyaline cells of textura angularis or textura globosa to
textura prismatica. Paraphyses numerous, filiform, septate,
unbranched, evanescent. Asci unitunicate, cylindrical, long
pedicellate, apex obtuse, with J- apical ring. Ascospores
8-spored, ellipsoidal to subglobose, pale brown to dark
brown, 0–1-septate, smooth-walled, with guttules and a
germ pore at one or each end, without gelatinous append-
ages. Asexual morph: Undetermined (adapted from Barr
1993; Huhndorf etal. 1999a).
Type genus: Jobellisia M.E. Barr 1993
Notes: Jobellisia was established based on Letendraea
luteola as a member of Clypeosphaeriaceae (Barr 1993).
Subsequently, other species of Jobellisia which have been
found in aquatic and/or terrestrial habitats in Asia, Europe
and the USA, were reported as resembling J. luteola, in hav-
ing ascomata with necks and brown, 1-septate ascospores
with germ pore(s) (Barr 1994; Huhndorf et al. 1999a;
Ranghoo etal. 2001; Leroy 2006; Untereiner etal. 2013).
Huhndorf etal. (2004b) analyzed LSU sequence data for
J. fraterna and J. luteola and found that they located in
Diaporthales. However, Réblová (2008) transferred J. rhyn-
chostoma to Bellojisia (Lasiosphaeriaceae) and established
Jobellisiaceae as an independent family in Sordariomycetes
based on LSU sequence data. Liu etal. (2012) introduced
J. guangdongensis and found that Jobellisiaceae is basal
to Calosphaeriales and Diaporthales. Maharachchikum-
bura etal. (2015, 2016) introduced the monotypic Jobel-
lisiales, which is closely related to Calosphaeriales based
on the multi-gene analysis. Hyde etal. (2017) found that
Jobellisiales is sister to Togniniales with a divergence of ca.
130–188 MYA in the MCC tree. Hongsanan etal. (2017)
and Wijayawardene etal. (2018) suggested that Jobellisiales
should be reduced as a family in Calosphaeriales. Hyde etal.
(2020) and Wijayawardene etal. (2020) listed Jobellisiales
in Hypocreomycetidae. However, species of Jobellisiales
have cylindrical asci with rounded apices and ellipsoidal,
brown, 1-septate ascospores similar to those of Diaporthales
(Maharachchikumbura etal. 2016; Hyde etal. 2020). The
clade characterising Jobellisiales is phylogenetically distinct
from other orders of Diaporthomycetidae (Fig.1).
Jobellisia M.E. Barr, Mycotaxon 46: 60 (1993)
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, gregarious, semi-immersed to superficial, obpyriform
to subglobose, collapsing when dry or not, blue-green, yel-
low, brown to black, rough, ostiolate, with beaked to conical
necks, periphysate. Peridium membranaceous, comprising
Fungal Diversity
1 3
three layers, outer layer composed of brown or blue-green
or orange cells of textura angularis-globosa; middle layer
composed of brown to yellow cells of textura angularis;
inner layer composed of hyaline cells of textura prismatica.
Paraphyses numerous, filiform, septate, unbranched, eva-
nescent. Asci unitunicate, cylindrical, long pedicellate, apex
obtuse, with J- apical ring. Ascospores 8-spored, ellipsoidal
to subglobose, pale brown to dark brown, straight to slightly
curved, 0–1-septate, smooth-walled, with guttules and a
germ pore at one or each end, without gelatinous append-
ages. Asexual morph: Undetermined (adapted from Barr
1993; Huhndorf etal. 1999a).
Notes: Barr (1993) established Jobellisia based on
Letendraea luteola. Eight species have been accommo-
dated in Jobellisia; however, only three (Jobellisia fraterna
SMH2863, J. guangdongensis HMAS 251240 and J. luteola
SMH2753) have molecular data (Huhndorf etal. 1999a;
Réblová 2008; Liu etal. 2012). More phylogenetic infor-
mation is needed to determine their taxonomic placement.
Type species: Jobellisia luteola (Ellis & Everh.) M.E. Barr,
Mycotaxon 46: 61 (1993)
Basionym: Letendraea luteola Ellis & Everh., Proc. Acad.
nat. Sci. Philad. 47: 415 (1895)
Facesoffungi number: FoF 10000; Fig.8
Saprobic on wood. Sexual morph: Ascostromata crus-
tose. Ascomata 460–570 × 515–620µm (x̄ = 500 × 560µm,
n = 10), perithecial, gregarious, superficial, obpyriform
to subglobose, collapsing when dry, black to dark brown,
rough, ostiolate, with conical necks, periphysate. Perid-
ium 40–75µm (x̄ = 55µm, n = 30) wide, membranaceous,
comprising three layers, outer layer composed of brown to
orange cells of textura angularis-globosa; middle layer com-
posed of brown to pale brown cells of textura angularis;
inner layer composed of hyaline cells of textura prismatica.
Paraphyses numerous, filiform, septate, unbranched, eva-
nescent. Asci unitunicate, cylindrical, long pedicellate, apex
obtuse, with J- apical ring. Ascospores 12–14 × 4.5–5.5µm
(x̄ = 13.5 × 5µm, n = 50), 8-spored, ellipsoidal to subglobose,
pale brown to dark brown, straight to slightly curved, 1-sep-
tate, smooth-walled, with guttules and a germ pore at each
end, without gelatinous appendages. Asexual morph: Unde-
termined (adapted from Barr 1993; Huhndorf etal. 1999a).
Material examined: USA, Michigan, Marquette, Huron
Mountain Club, end of road at Mountain Lake, 45º 00′ N/87°
0′ W, on dead wood over the water, 17 August 1997, S.M.
Huhndorf (F-SMH 3349).
Known hosts and distribution: On decayed wood in Ohio,
USA (type locality) (Ellis and Everhart 1895); on decayed
decorticated wood in Iowa, Michigan, North Carolina, New
York and Ontario, USA (Barr 1993; Huhndorf etal. 1999a).
Notes: Jobellisia luteola has been collected in tem-
perate regions (Barr 1993; Huhndorf et al. 1999a).
Maharachchikumbura etal. (2016) re-examined the holo-
type (NY00914363) collected by J.P. Morgan in Ohio,
USA. Huhndorf etal. (1999a) also introduced two new
specimens of J. luteola, SMH 2753 and SMH 3349. Sub-
sequently, Huhndorf etal. (2004b) analyzed LSU sequence
data for SMH 2753 which is widely used to represent J.
luteola. In this study, the authentic material of SMH 3349
was re-examined.
Other genera inDiaporthomycetidae
Ophiostomatales Benny & Kimbr.
Ophiostomataceae Nannf.
Fragosphaeria Shear, Mycologia 15(3): 124 (1923)
Saprobic on wood. Sexual morph: Ascomata solitary,
superficial, carbonaceous or coriaceous, globose, black,
glabrous or surrounded by hyaline hyphae. Asci 8-spored,
unitunicate, globose. Ascospores ovoid to reniform, asep-
tate, guttulate, smooth-walled. Asexual morph: Hyphomy-
cetous. Conidiophores hyaline, branched. Conidia hyaline,
aseptate, suballantoid to ellipsoidal (adapted from Saccardo
1881; Shear 1923).
Notes: Fragosphaeria is typified by F. purpurea and
has cleistothecial ascomata, globose asci and reniform
ascospores (Shear 1923). Chesters (1935) transferred this
genus to Cephalotheca based on similar morphologies. In
this study, we find that the sexual characteristics of Fra-
gosphaeria are similar to members of Eurotiales (Euroti-
omycetes), but the asexual morph is unknown. However,
Suh and Blackwell (1999) found that F. purpurea is closely
related to Ophiostoma (Ophiostomatales) based on LSU and
SSU sequence analyses. Hence, Fragosphaeria is accepted
as a genus in Ophiostomataceae (Diaporthomycetidae)
(Hyde etal. 2020).
Type species: Fragosphaeria purpurea Shear, Mycologia
15(3): 124 (1923)
Facesoffungi number: FoF 10001; Fig.9
Saprobic on wood. Sexual morph: Ascomata 90–130µm
(x̄ = 120µm, n = 10) diam., cleistothecial, solitary, superfi-
cial, carbonaceous, globose, dark brown to black. Peridium
brittle, composed of carbonaceous, brown to reddish brown
cells of textura prismatica, squashed fragments pentagonal.
Asci 4–6µm (x̄ = 5µm, n = 20) diam., 8-spored, unitunicate,
globose. Ascospores 2.5–3 × 1.5–2.8µm (x̄ = 2.8 × 2µm,
n = 20), overlapping, reniform, aseptate, hyaline when
young, becoming pale brown to olivaceous when mature,
guttulate, smooth-walled. Asexual morph: Hyphomycetous.
Conidiophores hyaline, branched. Conidia hyaline, aseptate,
suballantoid to ellipsoidal (adapted from Shear 1923).
Material examined: USA, Virginia, on culture of Pila-
cre sp., July 1920, C.L. Shear (BPI-566225, type); Swe-
den, Göteborg, Nature Park of Botanical Garden, on
Fungal Diversity
1 3
Fig. 8 Jobellisia luteola: a–l (F-SMH 3349); m, n (redrawn from
Huhndorf et al. 1999a). a Herbarium material label. b Collapsing
ascomata. c Ascomata on host. d Ascoma in cross section. e Perid-
ium. f Ostiole with periphyses. g–l Ascospores (arrow indicates a
germ pore). m Ascospores and asci with apical thickening and long
pedicel. n Septate paraphyses. Scale bars: c = 500µm, d = 200µm, f,
n = 50µm, e, m = 20µm, g–l = 10µm
Fungal Diversity
1 3
dead deciduous wood, 15 October 1970, U. Eliasson
(BPI-566226).
Known hosts and distribution: On culture of Pilacre
petersii in USA (type locality) (Shear 1923); on dead decid-
uous wood in UK, Sweden (Eliasson 1971; Yaguchi etal.
2006).
Notes: Shear (1923) introduced Fragosphaeria purpurea,
which was found in the culture of Pilacre petersii in Vir-
ginia, September 1920, as the generic type. In this study,
we re-examined type material (566,225), which was iso-
lated from a culture of Pilacre collected in Virginia, July
1920. This specimen is a dry fragmented culture, and we
were unable to find any structure for observation. We also
obtained authentic material, 566,226 collected and deter-
mined by Eliasson (1971).
Genera transferred toHypocreomycetidae
Coronophorales genera incertae sedis
Tengiomyces Réblová, Mycotaxon 70: 408 (1999)
Saprobic on wood. Sexual morph: Ascomata perithecial,
superficial, solitary to gregarious, globose to subglobose,
coriaceous, dark brown to black, surrounded by setae, seated
on sparse, brown, septate, branched hyphae, ostiolate, with
papilla. Setae brown to dark brown, spine-like, straight,
acute, aseptate, opaque, unbranched. Peridium membrana-
ceous, composed of brown to hyaline cells of textura angu-
laris to textura prismatica. Paraphyses septate, evanescent.
Asci 8-spored, unitunicate, cylindrical to clavate, broadly
rounded to truncate and thickened at the apex, pedicellate.
Ascospores ellipsoidal to clavate, with broadly rounded
Fig. 9 Fragosphaeria purpurea: (BPI-566226). a Herbarium material label. b Ascoma. c, d Squash mounts of ascoma (d in cotton blue). e–g
Asci (g in Melzer’s reagent). h Ascospores. Scale bars: b = 100µm, c = 50µm, d = 20µm, e–h = 5µm
Fungal Diversity
1 3
ends, straight to slightly curved, septate, middle cell(s) pale
brown to brown, end cell(s) hyaline to subhyaline, slightly
constricted at the septa, smooth-walled. Asexual morph
(associated): Hyphomycetous, spadicoides-like. Mycelium
semi-immersed to superficial. Conidiophores arising from
superficial hyphae growing from the base of the perithecial
initials and mature ascomata, macronermatous, mononema-
tous, unbranched or branched in the lower parts, cylindrical,
pale brown, straight, flexuous in the lower parts, septate, not
constricted or slightly constricted at the septa, apical cells
rounded. Conidiogenous cells polytretic, integrated, inter-
calary and terminal. Conidia obpyriform to clavate, broadly
rounded at the apex, septate, not constricted or slightly con-
stricted at the septa, brown or the pigment fading from apex
to bottom (adapted from Réblová 1999a).
Notes: Réblová (1999a) transferred Chaetosphaerella
indica to Tengiomyces and established a monotypic genus.
The black ascomata with setae on T. indicus are simi-
lar to Helminthosphaeria and Chaetosphaerella, and the
ascospores comprise two brown middle cells and one hya-
line cell at each end and are similar to Crassochaeta and
Chaetosphaerella (Chaetosphaerellaceae, Coronophorales)
(Réblová 1999a; Huhndorf etal. 2004a). A hyphomycetous
spadicoides-like form was associated with T. indicus on the
natural substrate (Réblová 1999a). Spadicoides is polyphy-
letic in Pleosporales (Dothideomycetes) (Ma etal. 2016;
Réblová etal. 2018). However, we could not determine the
relationship between Tengiomyces and Spadicoides without
appropriate cultural characteristics and phylogenetic analy-
sis. In this study, Tengiomyces with black, bristly ascomata
and ellipsoidal ascospores comprise brown and hyaline cells
is similar to Chaetosphaerella rather than Helminthospha-
eria. Thus, we place Tengiomyces in Coronophorales genera
incertae sedis pending further studies with molecular data.
Type species: Tengiomyces indicus (Varghese & V.G.
Rao) Réblová, Mycotaxon 70: 408 (1999)
Basionym: Chaetosphaerella indica Varghese & V.G.
Rao, Biovigyanam 5(1): 2 (1979)
Facesoffungi number: FoF 10002; Fig.10
Saprobic on wood. Sexual morph: Ascomata
200–250µm (x̄ = 230µm, n = 5) diam., perithecial, super-
ficial, solitary to gregarious, globose to subglobose, cori-
aceous, dark brown to black, surrounded by setae, seated
on sparse, brown, septate, branched hyphae 3–8µm wide,
ostiolate, with papilla. Setae 3.5–11µm wide, brown to
dark brown, spine-like, straight, acute, aseptate, opaque,
unbranched. Peridium 20–26µm wide, comprising two lay-
ers, outer layer composed of brown cells of textura angu-
laris; inner layer membranaceous, composed of hyaline
cells of textura prismatica. Paraphyses septate, evanescent.
Asci 60–70 × 10–15µm, 8-spored, unitunicate, cylindrical
to clavate, broadly rounded to truncate and thickened at
the apex, with short pedicel. Ascospores 10–15 × 4–5.5µm
(x̄ = 13 × 5µm, n = 20), bi-seriate, ellipsoidal to clavate, with
broadly rounded ends, straight to slightly curved, 3-septate,
two middle cells pale brown to brown, two end cells hya-
line to subhyaline, slightly constricted at the septa, smooth-
walled. Asexual morph (associated): Hyphomycetous,
spadicoides-like. Mycelium semi-immersed to superficial.
Conidiophores 3.5–8µm wide, arising from superficial
hyphae growing from the base of the perithecial initials
and mature ascomata, macronematous, mononematous,
unbranched or branched in the lower parts, cylindrical, pale
brown, straight, flexuous in the lower parts, septate, not
constricted or slightly constricted at the septa, apical cells
rounded. Conidiogenous cells 4–6 × 5–6µm, polytretic, inte-
grated, intercalary and terminal. Conidia 14–18 × 5.5–10µm
(x̄ = 15 × 7.5µm, n = 20), obpyriform to clavate, broadly
rounded at the apex, 2–3-septate, not constricted or slightly
constricted at the septa, brown or the pigment fades from
apex to bottom (adapted from Réblová 1999a).
Material examined: China, An-Hui, LangYa Shan, on
dead wood of a deciduous tree, 20 June 1933, S.Q. Deng
(BPI-622100); Guiana, Saint Laurent, Piste Balate, 12km
from Saint Laurent, 20m elev, on dead branch, A. Rossman,
C. Feuillet and L. Skog, 19 November 1986 (BPI-622098).
Known hosts and distribution: On dead stems of a dicoty-
ledoneous plant in India (type locality) (Varghese and Rao
1979); on dead wood of a deciduous tree in China (Réblová
1999a); on dead branch in Guiana.
Notes: Réblová (1999a) introduced Tengiomyces indicus
after examining the holotype of Chaetosphaerella indica
(AMH 3871), collected by Varghese in India (Varghese and
Rao 1979) and a sample BPI 622100 collected by Deng in
China. We re-examined two authentic specimens (622100
and 622098) which were determined by Réblová (mentioned
in the label of material) and found that they were dry and
observed only ascomata and evanescent asci and ascospores.
We also found a spadicoides-like taxon by T. indicus on the
natural substrate (Fig.10h–i).
Hypocreales genera incertae sedis
Sulcatistroma A.W. Ramaley, Mycotaxon 93: 140 (2005)
Saprobic on leaves. Sexual morph: Ascostromata dis-
crete, multiloculate, erumpent through leaf epidermis,
ellipsoidal, rarely circular. Ascomata perithecial, immersed,
formed in stroma, subglobose, dark brown, ostiole with
hyaline periphyses. Peridium membranaceous, composed
of hyaline to brown cells of textura angularis. Paraphy-
ses numerous, septate, slightly constricted at the septa,
unbranched, tapering to a rounded apex. Asci 8-spored, uni-
tunicate, clavate, short pedicellate. Ascospores bi-seriate,
hyaline, allantoid with rounded ends, aseptate, smooth-
walled. Asexual morph: Hyphomycetous. Conidiophores
arising from aerial hyphae, erect, inflated at the base. Phi-
alides monophialidic, smooth-walled, hyaline, cylindrical,
Fungal Diversity
1 3
unbranched or with sympodial conidiogenous cells. Conidia
hyaline to pale orange, ellipsoidal to allantoid, aseptate,
smooth-walled, with rounded ends (adapted from Ramaley
2005).
Notes: The monotypic genus Sulcatistroma has ellipsoidal
ascostromata, clavate asci, allantoid ascospores and ellip-
soidal to allantoid conidia (Ramaley 2005). This genus is
similar to the sexual morph of Diatrype (Xylariales), dif-
fering from Sulcatistroma in having sympodial conidiog-
enous cells, whereas Diatrype has percurrently proliferating
conidiogenous cells (Ramaley 2005). Initially, Sulcatistroma
was accepted as a member of Calosphaeriales based on its
spicate clusters of asci (Ramaley 2005), and later it was
included in Calosphaeriaceae (Lumbsch and Huhndorf 2010;
Maharachchikumbura etal. 2015, 2016; Wijayawardene
etal. 2018). Vu etal. (2019) analyzed ITS sequence data for
ex-type strain Sulcatistroma nolinae (CBS 117709). This
strain is more closely related to members of Nectriaceae
(Hypocreales) (Fig.1). Sulcatistroma has similar features to
Nectriaceae species, such as clavate asci, aseptate, allantoid
Fig. 10 Tengiomyces indicus:
a–c (BPI-622098); d–f, h, i
(BPI-622100); g, j (redrawn
from Réblová 1999a). a Her-
barium material. b–d Ascomata
on host. e Squashed ascoma
with dark brown setae. f Ascus.
g Ascus with ascospores. h
Conidiophore with conidium. i
Conidium. j Conidiophores of
spadicoides-like from nature.
Scale bars: b, d = 500µm,
c = 200µm, e = 100µm, f–h,
j = 20µm, i = 10µm
Fungal Diversity
1 3
ascospores and phialophora-like asexual morphs, but does
not have the red ascomata which is characteristic of Nec-
triaceae (Hirooka etal. 2012). We place Sulcatistroma in
Hypocreales genera incertae sedis pending phylogenetic
data.
Type species: Sulcatistroma nolinae A.W. Ramaley,
Mycotaxon 93: 140 (2005)
Facesoffungi number: FoF 10003; Fig.11
Saprobic on leaves. Sexual morph: Ascostromata dis-
crete, multiloculate, erumpent through leaf epidermis, ellip-
soidal, rarely circular, with long axis parallel to length of
leaf, 900–1500 × 350–850µm (x̄ = 1200 × 450µm, n = 5).
Ascomata 120–250µm diam., perithecial, immersed, sub-
globose, brown to dark brown, ostiole with hyaline peri-
physes. Peridium 7–25µm (x̄ = 16.5 µm, n = 20) wide,
membranaceous, composed of hyaline to pale brown
cells of textura angularis. Paraphyses numerous, septate,
slightly constricted at the septa, unbranched, tapering to
the rounded apex. Asci 8-spored, unitunicate, clavate, short
pedicellate, apical ring not apparent. Ascospores bi-seriate,
hyaline, allantoid with rounded ends, aseptate, smooth-
walled. Asexual morph: Hyphomycetous, phialophora-like.
Fig. 11 Sulcatistroma nolinae:
b–g (BPI-864276, holotype);
a. h, i (redrawn from Ramaley
2005). a Conidiophores with
conidia. b Herbarium material.
c Immersed ascostroma. d Mul-
tiloculate ascostroma. e Ascoma
in cross section. f Peridium. g
Filament-like support struc-
tures in ascoma. h Asci and
ascospores. i Tapering and
septate paraphyses. Scale bars:
a, h, i = 20µm, c = 200µm,
d = 100µm, e–g = 50µm
Fungal Diversity
1 3
Conidiophores arising from aerial hyphae, erect, inflated at
the base. Phialides monophialidic, smooth-walled, hyaline,
cylindrical, unbranched or an axis with 1–4 sympodial con-
idiogenous cells. Conidia hyaline to pale orange, ellipsoi-
dal to allantoid, aseptate, smooth-walled, with rounded ends
(adapted from Ramaley 2005).
Material examined: USA, New Mexico, Lincoln Co., Val-
ley of Fires, roadside, on leaves of Nolina micrantha, A.W.
Ramaley, 9 October 2002 (BPI-864276, holotype).
Known hosts and distribution: On decayed leaves of
Nolina micrantha in the USA (type locality) (Ramaley 2005).
Notes: We re-examined the holotype (864,276), but the
ascomata contained only some filament-like supporting
structures (Fig.11e–g). Therefore, we provided hand draw-
ings of other characteristics (Fig.11a, h–i) based on Rama-
ley (2005).
Sordariomycetidae O.E. Erikss. & Winka
The subclass Sordariomycetidae was introduced by Eriks-
son and Winka (1997) and previously comprised six orders.
Eight orders with 23 families are now included based on phy-
logenetic and morphological studies (Maharachchikumbura
etal. 2015; Hongsanan etal. 2017; Hyde etal. 2017, 2020;
Marin-Felix etal. 2020; Wijayawardene etal. 2020). Sor-
dariomycetidae members are widespread and usually as
saprobes, pathogens or endophytes (Muggia etal. 2015;
Maharachchikumbura etal. 2016; Hyde etal. 2020). Fig-
ure1 reveals that this subclass is sister to Diaporthomyceti-
dae (68%ML) in Sordariomycetes (Fig.12).
Boliniales P.F. Cannon, in Kirk etal., Dictionary of Fungi
9 (2001)
The saprotrophic Boliniales was introduced by Kirk
etal. (2001) and is characterized by black ascomata,
cylindrical asci and fusiform to subglobose ascospores
(Maharachchikumbura etal. 2015, 2016; Hyde etal.
2020). Some members of this order with carbonaceous
ascomata were traditionally linked to Xylariales, e.g.
Camarops (Karsten 1873). Boliniales was clarified by
Maharachchikumbura etal. (2015) and currently contains
nine genera based on morphology and phylogenetic analy-
ses (Hyde etal. 2020; Wijayawardene etal. 2020). This
order is sister to Phyllachorales (Maharachchikumbura
etal. 2016; Hongsanan etal. 2017; Hyde etal. 2017; this
study, 83%ML, Fig.1).
0.05
Apiorhynchostoma apiosporum UAMH 11088
Camarops ustulinoides SMH1988
Camarops rogersii SMH3001
Camarops amorpha SMH1450
Phialemonium inflatum CBS:259.39
Cornipulvina ellipsoides SMH1378
Camarops microspora CBS 649.92
Camaropella lutea ANM936
Camarops petersii JM1655
Phialemonium atrogriseum CBS:604.67
Endoxyla macrostoma UAMH 11491
Camarops polysperma 2006-747
Endoxyla operculata UAMH 11085
Phialemonium obovatum CBS:279.76
Camaropella pugillus CBS 128346
Camarops plana Pouzar s.n.
Camarops scleroderma MP 4464
Endoxyla mallochii UAMH 11087
Camarops tubulina SMH4614
Pseudovalsaria ferruginea UAMH 11490
8 6/0.97
9 5/0.99
98/--
100/0.97
89/0.93
100/1.00
8 4/--
71/0.93
7 2/--
Cephalothecales
Boliniales
Boliniaceae
6 8/--
7 0/--
53/--
50/--
Camarops
Camaropella
Cam. amorpha
Cam. microspora
Cam. plana
Apiorhynchostoma
Pseudovalsaria
Cornipulvina
Endoxyla
Fig. 12 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS and RPB2 sequence data of Boliniales.
The confidence values of bootstrap (BS) proportions from the Maxi-
mum 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 lack of statistical support (< 50% for ML-BS and < 0.90
for BY-PP). Twenty strains are included in the combined analyses
which comprise 2485 characters (960 characters for LSU, 475 char-
acters for ITS, 1050 characters for RPB2) after alignment. Strains
of Cephalothecales are used as the outgroup taxa. The best score in
IQ-TREE explores with a final likelihood value of -9890.4903 is pre-
sented. The model of each partitioned gene is: LSU: GTR + I + G;
ITS: GTR + G; RPB2: GTR + I + G. The strain numbers are noted
after the species names. Ex-type strains are in bold. Alignments are
available at TreeBASE (URL: http:// purl. org/ phylo/ treeb ase/ phylows/
study/TB2:S28264)
Fungal Diversity
1 3
Boliniaceae Rick, Brotéria, sér. bot. 25(2–3): 65 (1931)
Saprobic on wood in terrestrial habitats. Sexual morph:
Ascostromata immersed, erumpent or superficial, pale brown
to black, irregular in shape, surface furfuraceous or sur-
rounded by hyaline to brown hyphae. Ascomata perithecial,
solitary or gregarious, immersed, erumpent or superficial,
globose to obpyriform, brown to black, papillate. Peridium
carbonaceous or coriaceous or membranaceous, composed
of brown to dark brown cells of textura intricata or textura
prismatica. Paraphyses abundant, filamentous, tapering,
septate, branched. Asci 8-spored, unitunicate, cylindrical
to clavate, apex rounded to blunt, pedicellate. Ascospores
2–3-seriate, normally cells in the center brown to dark
brown, and the terminal cell(s) hyaline to pale brown, sub-
globose to ellipsoidal, 0–2-septate, smooth-walled, guttulate.
Asexual morph: Undetermined (adapted from Rick 1931;
Romero and Samuels 1991; Maharachchikumbura etal.
2016).
Type genus: Camarops P. Karst. 1873 (= Bolinia
(Nitschke) Sacc. 1882)
Notes: Boliniaceae, introduced by Rick (1931), is char-
acterized by black ascostromata and cylindrical asci with
smooth-walled ascospores. Earlier, this family was placed
in Xylariales based on morphological similarities, such as
carbonaceous ascomata (Barr 1990; Romero and Samuels
1991). Andersson etal. (1995) relied on analyzing SSU
sequence data and concluded that Boliniaceae was more
closely related to Sordariales. This family was subsequently
used to establish Boliniales by Kirk etal. (2001). Thus,
Boliniaceae, based on Bolinia is typified by B. tubulina, is
the only member of Boliniales (Maharachchikumbura etal.
2016; Kirk etal. 2001). Nannfeldt (1972) recommended
Bolinia should be synonymized as Camarops which is typi-
fied by Camarops hypoxyloides (Karsten 1873) as they have
similar characters. Phylogenetically, Boliniales is closely
related to Phyllachorales and Sordariales in Sordariomy-
cetidae (Huhndorf etal. 2004b; Miller and Huhndorf 2005;
Hongsanan etal. 2017; Hyde etal. 2017; this study, Fig.1).
However, since both Boliniales and Boliniaceae were estab-
lished based on Bolinia tubulina, even if B. tubulina was
transferred to Camarops tubulina, this species is still the
type of its order and family (Maharachchikumbura etal.
2016). We accept Camarops tubulina as type species of
Boliniales and Boliniaceae, and Camarops hypoxyloides
as the type of Camarops, as established (Karsten 1873;
Rick 1931; Kirk etal. 2001); and more characteristics and
phylogenetic information are expected to determine the
affinities of Camarops and other genera in the Boliniaceae.
Nine genera are accepted into this family, six of which have
available molecular data (viz. Apiorhynchostoma, Camaro-
pella, Camarops, Cornipulvina, Endoxyla, Pseudovalsaria)
(Huhndorf and Miller 2008; Raja etal. 2011; Untereiner
etal. 2013; Maharachchikumbura etal. 2015, 2016; Vu etal.
2019; Hyde etal. 2020; Wijayawardene etal. 2020), and Api-
ocamarops, Mollicamarops, Neohypodiscus were included
because of their morphology similar to Camarops (Anders-
son etal. 1995; Rogers and Ju 2003; Vasilyeva 2007). In
this study, we find that some species of ascospores are not
arranged in a uniform direction in the ascus. For example,
in Apiocamarops, ascospore is composed of a large brown
cell and a small hyaline cell (Samuels and Rogers 1987).
Ascospores partly are oriented with the hyaline cell to the
ascus apex; and partly in the opposite direction to the ascus
base.
Apiocamarops Samuels & J.D. Rogers, Mycotaxon 28(1):
54 (1987)
Saprobic on wood. Sexual morph: Ascostromata super-
ficial or semi-immersed, brown to black, irregular in shape,
surface furfuraceous or glabrous. Ascomata perithecial,
gregarious, superficial or semi-immersed, globose to sub-
globose, membranaceous, orange, brown to black, ostiolate,
with periphyses. Peridium membranaceous, comprising two
layers, outer layer composed of brown cells of textura angu-
laris; inner layer composed of hyaline cells of textura pris-
matica. Asci 8-spored, unitunicate, cylindrical, pedicellate,
apex rounded, with J- apical ring. Ascospores uniseriate,
0–1-septate, unequal, oblong to ellipsoidal, smooth-walled,
hyaline when young, composed of a large, brown cell with
or without a terminal pore and a small, hyaline cell when
mature, guttulate. Asexual morph: Undetermined (adapted
from Samuels and Rogers 1987; Rogers 1988; Reagan and
Waide 1996; Rogers and Ju 2003).
Notes: Apiocamarops was established by Samuels and
Rogers (1987) and resembles Camarops. Both genera have
similar cylindrical asci and ellipsoidal ascospores with germ
pore(s), but the ascospores in Apiocamarops are 1-septate
and apicosporous, while those of Camarops lack septum
(Samuels and Rogers 1987; Rogers 1988; Reagan and Waide
1996; Rogers and Ju 2003). Four species are accommodated
in this genus, but molecular data is lacking.
Type species: Apiocamarops alba Samuels & J.D. Rog-
ers, Mycotaxon 28(1): 54 (1987)
Facesoffungi number: FoF 10004; Fig.13
Saprobic on decorticated wood. Sexual morph: Ascos-
tromata superficial, brown to black, irregular in shape,
surface furfuraceous. Ascomata 450–750 × 350–700 µm
(x̄ = 550 × 450µm, n = 10), per ithecial, gregarious, super-
ficial, globose to subglobose, brown to black, papillate,
ostiole central, with periphyses, covered by white to yel-
lowish, furfuraceous layer. Peridium 35–60µm (x̄ = 45µm,
n = 30) wide, membranaceous, comprising two layers,
outer layer composed of dark brown to pale brown cells
of textura angularis; inner layer composed of hyaline
cells of textura prismatica. Paraphyses 1–3µm wide,
numerous, filiform, septate, branched, longer than the
Fungal Diversity
1 3
asci. Asci 75–95(–110) × 4–6 µm (x̄ = 90 × 5µm, n = 50),
8-spored, unitunicate, cylindrical, pedicel short, apex
rounded, with J-inconspicuous apical ring. Ascospores
(4.5–)5.5–6.5(–8) × 1.5–3µm (x̄ = 6 × 2.5µm, n = 50), uni-
seriate, oblong to ellipsoidal, slightly curved, 1-septate,
unequal, hyaline when young, composed of a large, brown
cell with a minute terminal pore and a small, hyaline cell
when mature, partly hyaline cell towards the ascus apex;
partly in the opposite direction, smooth-walled, guttulate.
Asexual morph: Undetermined.
Material examined: Brazil, Amazonas, Plateau of
Serra Araca, N side of North Mountain, alt. 1250m.,
00° 57' N, 63° 21' W, cloud forest, on decorticated wood,
19–24 February 1984, G.J. Samuels, G.T. Prance and J.
Pipoly (NY-00986029, holotype); ibid. (NY-00986030,
paratype).
Fig. 13 Apiocamarops alba: a,
c–e, h, l, m, o (NY-00986029,
holotype); b, f, g, i–k, n, p
(NY-00986030, paratype).
a Herbarium material. b
Ascostroma on host. c Papilla
without furfuraceous cover. d,
e Ascomata. f, g Ascomata in
cross section (f from dry slide).
h Peridium. i Paraphyses. j–l
Asci. m–p Ascospores (o arrow
indicates terminal pore). Scale
bars: b = 2mm, c. g = 200µm,
d. e = 500µm, f = 100µm,
h–i = 50µm, j–l = 20µm,
m–p = 5µm
Fungal Diversity
1 3
Known hosts and distribution: On decorticated wood in
Brazil (type locality), Guiana, Venezuela (Samuels and Rog-
ers 1987).
Notes: As noted by Samuels and Rogers (1987), we re-
examined the type materials, 00986029 and 00986030,
which are well preserved.
Apiorhynchostoma Petr., Annls mycol. 21(3/4): 185 (1923)
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, scattered, immersed to semi-immersed, globose to
subglobose, dark brown to black, papillate, ostiole central,
with periphyses. Peridium membranaceous, comprising two
layers, outer layer composed of brown to pale brown cells
of textura angularis; inner layer composed of hyaline cells
of textura prismatica. Asci 8-spored, unitunicate, cylindri-
cal, pedicellate, with J- apical ring. Ascospores uniseriate,
oblong to ellipsoidal, straight to slightly curved, 0–2-septate,
hyaline when young, becoming composed of 1 or 2 large,
brown cell(s) with a minute pore at the end, other end with
a small, hyaline to pale brown cell, smooth-walled or ver-
rucose, guttulate. Asexual morph: Undetermined (adapted
from Petrak 1923).
Notes: Currey (1859) introduced two species, named
‘Sphaeria apiculata’ and ‘Sphaeria curreyi Blox. MSS.’.
Rabenhorst (1860) examined some materials and stated that
Sphaeria apiculata Curr. is illegitimate as it has a homo-
nym of Sphaeria apiculata Wallr. (Wallroth 1833). Thus,
Rabenhorst (1860) re-named S. apiculata Curr. as Sphaeria
curreyi Rabenh. which is a homonym of S. curreyi Blox.
MSS. Due to the illegitimate species naming of ‘Sphaeria
apiculata’ and ‘Sphaeria curreyi’, subsequent research has
been controversial. Petrak (1923) established Apiorhynchos-
toma and named S. apiculata Curr. as Apiorhynchostoma
apiculatum as the type species. Winter (1886) and Müller
and von Arx (1962) recognized the former species related
to ‘Sphaeria curreyi’ and transferred S. curreyi Rabenh.
as Apiorhynchostoma curreyi as the type species of Api-
orhynchostoma (Rogers etal. 1994a; Hyde etal. 1998;
Stadler etal. 2013; Untereiner etal. 2013). In our study,
we proposed a new epithet, Apiorhynchostoma apiosporum,
based on isotype material of Sphaeria curreyi Rabenh. to
avoid future confusion. Therefore, the illegitimate A. api-
culatum (≡ S. apiculata Curr.) and A. curreyi (≡ S. curreyi
Rabenh.) are replaced by A. apiosporum. Apiorhynchostoma
is characterized by subglobose ascomata and ascospores
which are composed of two brown cells and a hyaline
terminal cell (Müller and von Arx 1962; Rappaz 1995;
Untereiner etal. 2013). This genus has been included in
Amphisphaeriaceae and Clypeosphaeriaceae based on their
similar ascomata and ascospores (Hyde etal. 1998; Réblová
1998; Wang etal. 2004). Untereiner etal. (2013) analyzed
ITS-LSU sequence data for ‘A. curreyi’ and showed that
Apiorhynchostoma is closely related to Cornipulvina and
Pseudovalsaria in Boliniales. In this study, the strain name
of A. curreyi is replaced by Apiorhynchostoma apiosporum
and it is sister to Pseudovalsaria ferruginea in Boliniales
(72%ML, Fig.12).
Type species: Apiorhynchostoma apiosporum S.K.
Huang & K.D.Hyde, nom. nov.
Synonym: Sphaeria apiculata Curr., Trans. Linn. Soc.
London 22: 326 (1859)
Sphaeria curreyi Rabenh., Fungi europ. exsicc. Klotzschii
herbarii vivi mycologici continuatio, Edn nova. Series
secunda, Cent. 3: no. 250 (1860)
Apiorhynchostoma apiculatum (Sacc.) Petr. [as 'apicu-
lata'], Annls mycol. 21(3/4): 185 (1923)
Apiorhynchostoma curreyi E. Müll., in Müller & von Arx,
Beitr. Kryptfl. Schweiz 11(no. 2): 706 (1962)
Facesoffungi number: FoF 10005; Index Fungorum num-
ber: IF558201; Fig.14
Etymology: apiosporum refers to the ellipsoidal
ascospores with one hyaline cell markedly smaller than the
other.
Saprobic on decorticated wood. Sexual morph: Asco-
mata 530–660 × 470–580 µm (x̄ = 590 × 530µm, n = 10),
perithecial, scattered, immersed to semi-immersed, glo-
bose to subglobose, dark brown to black, papillate, ostiole
central, with periphyses. Peridium 15–65µm (x̄ = 35µm,
n = 30) wide, membranaceous, 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 2–4.5µm wide, numerous, filiform,
septate, longer than asci. Asci 190–200(–250) × 7.5–13µm
(x̄ = 198 × 10µm, n = 50), 8-spored, unitunicate, cylindrical,
pedicellate, apex rounded, with J- inconspicuous apical ring.
Ascospores 23.5–30 × 5.5–9µm (x̄ = 25 × 7.5 µm, n = 50),
uniseriate, oblong to ellipsoidal, straight to slightly curved,
1–2-septate, unequal, hyaline when young, becoming com-
posed of 1 or 2 large, brown cell(s) with a minute pore and a
hyaline, thick-walled at the end, other end with a small, hya-
line to pale brown cell, smooth-walled, guttulate. Asexual
morph: Undetermined.
Material examined: UK, Batheaston, on dead decorti-
cated wood, F. Currey, Rabenhorst Fungi Europaei Exsiccati
nr. 250 (S-F61087, lectotype).
Known hosts and distribution: on decayed decorticated
wood of conifers in Austria, Canada, Germany, Sweden, UK
(type locality) (Currey 1859; Untereiner etal. 2013).
Notes: Currey (1859) described samples as being col-
lected on dry old wood of the South Western Railway, not far
from the Weybridge Station. Rabenhorst (1860) mentioned
that Currey collected material of Sphaeria curreyi Rabenh.
in Batheaston in 1856 and 1857, and this material was deter-
mined as the isotype of S. curreyi Rabenh. (Untereiner etal.
2013). As noted in S-F61087, this material was collected by
Currey in Batheaston and determined by Rabenhorst. Thus,
Fungal Diversity
1 3
we proposed this material as lectotype of Apiorhynchostoma
apiosporum.
Camaropella Lar.N. Vassiljeva, Mikol. Fitopatol. 31(1): 6
(1997)
Saprobic on wood. Sexual morph: Ascostromata erect,
immersed to semi-immersed, erumpent through bark of host,
carbonaceous, black, glabrous, pulvinate, peltate, discoid or
valsoid, hemispherical or spherical, sometimes with broad
attachment to the substrate. Ascomata perithecial, gregari-
ous, carbonaceous, cylindrical, black, ostiolate, with necks,
periphysate. Asci 8-spored, unitunicate, cylindrical, apex
rounded, pedicellate, with J- apical ring. Ascospores uni-
seriate, cylindrical to ellipsoidal, aseptate, pale brown or
dark brown, smooth-walled, with guttules. Asexual morph:
Undetermined (adapted from Vasilyeva 1997; Vasilyeva
etal. 2007).
Notes: Vasilyeva (1997) transferred Camarops pugillus (≡
Sphaeria pugillus Schw.) to Camaropella as generic type.
Camaropella species do not have a lopadostomoid type of
stromata as found in Camarops, but has special eutypoid
and eutypelloid or valsoid forms (Vasilyeva 1997). She also
introduced a second species, Camaropella lutea (Vasily-
eva etal. 2007). These two species are nested in Boliniales
Fig. 14 Apiorhynchostoma
apiosporum: (S-F61087,
lectotype). a Rotting, decor-
ticated wood. b Ascomata on
host. c Ascoma in cross section.
d Papilla with periphyses. e
Peridium. f Ascus with paraphy-
ses. g–j Asci. k, l Apical pore of
ascus (k arrow indicates pore).
m–r Ascospores (arrows indi-
cate the minute pore in n and a
hyaline, thick-walled at the end
in p–r). Scale bars: c = 200µm,
d, f–j = 50µm, e, k–r = 20µm
Fungal Diversity
1 3
and distant from Camarops tubulina based on phylogenetic
analysis (Huhndorf and Miller 2008; Untereiner etal. 2013;
Vu etal. 2019). In this study, strains of Camarops amorpha,
C. microspora and C. plana, which have pulvinate or peltate
stromata (Miller and Huhndorf 2005; Spatafora etal. 2006;
Raja etal. 2011), clustered with Camaropella species with
89%ML/0.93PP support (Fig.12), and this clade is distant
from Camarops clade (Untereiner etal. 2013; this study).
Therefore, we recommend transferring these three Cama-
rops species to Camaropella. However, Camaropella and
Camarops are very similar in morphology, in the cylindrical
asci, ellipsoidal ascospores and the arrangement type of the
ascomata in stroma. The relationship between Camarops and
Camaropella needs to be resolved with more collections and
sequence data.
Type species: Camaropella pugillus (Schwein.) Lar.N.
Vassiljeva, Mikol. Fitopatol. 31(1): 6 (1997)
Basionym: Sphaeria pugillus Schwein., Schr. naturf. Ges.
Leipzig 1: 38 [12 of repr.] (1822)
Facesoffungi number: FoF 10006; Fig.15
Saprobic on wood. Sexual morph: Ascostromata
2–3 × 1.5–3mm (x̄ = 2.2 × 2mm, n = 5) immersed to semi-
immersed, erumpent through bark of host, carbonaceous,
valsoid, black, glabrous. Ascomata 1.5–2.5 × 0.3–0.5mm
(x̄ = 1.8 × 0.4mm, n = 10), perit hecial, gregarious, cylin-
drical, black, ostiolate, with necks, circular in apical
Fig. 15 Camaropella pugil-
lus: (S-F24255). a Herbarium
material. b Semi-immersed
ascostroma. c Ascostroma
in longitudinal section. d
Ascostroma in apical view. e
Ascoma in cross section. f Neck
vertical section with periphyses.
g Peridium. h–j Asci (h arrow
indicates an inconspicuous
apical ring and i mounted in
Melzer's reagent). k Asci with
paraphyses. l–o Ascospores (n
arrow indicates germ pore at
end of ascospore and o mounted
in Melzer's reagent). Scale
bars: b, c = 2mm, d = 1mm,
e = 200µm, f = 100µm, g.
k = 50µm, h–j = 20µm,
l–o = 5µm
Fungal Diversity
1 3
view, periphysate. Peridium 25–60µm (x̄ = 50µm, n = 30)
wide, comprising two layers, outer layer carbonaceous,
composed of brown to black cells of textura angula-
ris; inner layer membranaceous, composed of hyaline
cells of textura prismatica. Paraphyses 1.5–3.5µm wide,
numerous, septate, unbranched, filiform, guttulate. Asci
(45–)47–53(–65) × 3.5–5.5µm (x̄ = 50 × 4.5 µm, n = 50),
8-spored, unitunicate, cylindrical, pedicellate, apex
rounded, with J- apical ring inconspicuous. Ascospores
4.5–6.5 × 2–4µm (x̄ = 6 × 3µm, n = 50), uniseriate, cylindri-
cal to ellipsoidal, aseptate, hyaline when young, becoming
pale brown or dark grey, sometimes slightly contracted at
middle of the cell, smooth-walled, 2 guttules, with a germ
pore at the base. Asexual morph: Undetermined.
Material examined: France, Aquitaine, Pyrénées-Atlan-
tiques, Pau 64, on dead wood of Quercus sp., 24 October
1998, F. Candoussau and G.J. Samuels (S-F24255); Sweden,
Uppland, Sollentuna, 700m NNW of Hägerstalund, on rotten
log, 3 May 1984, N. Lundqvist (UPS-UPS:BOT:F-120952).
Known hosts and distribution: On dead wood of Acer,
Liriodendron and Rhododendron in the USA (type locality)
(Shear 1940); on dead wood of Alnus glutinosa in Slovakia
(Vasilyeva 1997); on dead wood in Austria, Sweden, Rus-
sia (Vasilyeva 1997); on dead wood of Fraxinus excelsior
in Czech Republic (del Valle Catania and Romero 2003);
on decorticated wood of Podocarpus parlatorei in Argen-
tina (del Valle Catania and Romero 2003); on dead wood of
Quercus in France (del Valle Catania and Romero 2003).
Notes: We could not obtain the holotype, so we re-exam-
ined specimens F-120952 from UPS and F24255 from S her-
baria based on information provided by del Valle Catania and
Romero (2003). The strain of Camaropella pugillus nests in
Camaropella (Walker etal. 2012; Vu etal. 2019) and related
to Camaropella amorpha and Cam. microspora (Fig.12).
New combinations:
Camaropella amorpha (Boedijin) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Sarcostromella amorpha Boedijn, Persoonia
1(1): 17 (1959)
Synonym: Camarops amorpha (Boedijn) Nannf., Svensk
bot. Tidskr. 66(4): 362 (1972)
Index Fungorum number: IF558202
Camaropella microspora (P. Karst.) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Anthostoma microsporum P. Karst., Fungi
Fenniae Exsiccati, Fasc. 9: no. 860 (1869)
Synonym: Camarops microspora (P. Karst.) Shear, Myco-
logia 30(5): 588 (1938)
Index Fungorum number: IF558249
Camaropella plana (Pouzar) S.K. Huang & K.D. Hyde,
comb. nov.
Basionym: Camarops plana Pouzar, Česká Mykol. 40(4):
219 (1986)
Index Fungorum number: IF558250
Camarops P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 6, 53
(1873)
Synonym: Bolinia (Nitschke) Sacc., Syll. fung. (Abellini)
1: 352 (1882)
Saprobic on wood. Sexual morph: Ascostromata
immersed, erumpent to superficial, dark brown to black, car-
bonaceous or coriaceous, pulvinate, peltate, discoid or val-
soid, hemispherical, spherical or irregular, sometimes with
broad attachment to the substrate. Ascomata perithecial, gre-
garious, globose to subglobose, brown to dark brown, ostio-
late. Peridium comprising two layers, outer layer composed
of brown to dark brown cells of textura intricata; inner layer
composed of hyaline cells of textura prismatica. Paraphy-
ses abundant, persistent, filamentous, septate, branched. Asci
8-spored, unitunicate, cylindrical, pedicellate, apex rounded
to blunt, with J- apical ring. Ascospores 2–3-seriate, sub-
globose to ellipsoidal, aseptate, pale brown to dark brown,
smooth-walled, guttulate. Asexual morph: Undetermined
(adapted from Nannfeldt 1972).
Notes: Bolinia was introduced as a subgenus by Nitschke
(1867) and raised to generic rank by Saccardo (1882),
based on Sphaeria tubulina Alb. & Schw. Karsten (1873)
had earlier introduced Camarops based on C. hypoxyloides
P. Karst. Bolinia and Camarops, as very similar genera,
have often been discussed together in subsequent studies.
Munk (1957) and von Arx and Müller (1954) reviewed the
taxonomy of these taxa and concluded that Camarops and
Bolinia were distinct genera with similar characters, except
the former has monostichous ascomata (ascomata arranged
in one line or series in stroma, e.g. Figure16b) and the lat-
ter has polystichous (ascomata arranged in multiple lines
or groups in stroma, e.g. Figure17d–e). Shear (1938) and
Svrček (1969) recognized that Sphaeria tubulina Alb. &
Schw. should be a species of Camarops, named Camarops
tubulina. However, Nannfeldt (1972) re-examined species
of Camarops and Bolinia, and suggested Bolinia should be
merged with Camarops based on their similar characters.
Nannfeldt (1972) also proposed that the structure of the
ascospores, peridium and stroma should be emphasized in
the remodeled Camarops, and the arrangement of ascomata
in stroma and size, shape of stromata are considered to be
the basis for distinguishing species (del Valle Catania and
Romero 2003). Therefore, Bolinia was accepted as a syno-
nym of Camarops under Boliniaceae (Lumbsch and Huhn-
dorf 2010; Hyde etal. 2020; Wijayawardene etal. 2020).
Vasilyeva (1997) established Camaropella based on Cama-
rops pugillus. Untereiner etal. (2013) found that species
of Camaropella and Camarops sensu lato form a cluster
(see notes for Camaropella). In this study, we found that the
Fungal Diversity
1 3
previously proposed methods for distinguishing Camaro-
pella and Camarops morphologically are inconsistent with
phylogenetic analysis. Currently, we suggest using phylo-
genetic analysis as the main method to distinguish Cama-
ropella and Camarops. The complex needs resolving based
on more fresh collections.
Type species: Camarops hypoxyloides P. Karst., Bidr.
Känn. Finl. Nat. Folk 23: 53 (1873)
Facesoffungi number: FoF 10007; Fig.16
Saprobic on wood. Sexual morph: Ascostromata dark
brown to black, carbonaceous, pulvinate, discoid, hemi-
spherical or spherical, sometimes with broad attachment
to the substrate. Ascomata perithecial, gregarious, globose
to subglobose, brown to dark brown, monostichous, carbo-
naceous, ostiolate, with periphyses. Peridium 25–70µm
(x̄ = 40µm, n = 20) wide, comprising two layers, outer
layer carbonaceous to coriaceous, composed of brown to
dark brown cells of textura intricata to textura angularis;
inner layer membranaceous, composed of hyaline cells of
textura prismatica. Asci 8-spored, unitunicate, cylindri-
cal, apex rounded, pedicellate. Ascospores 6–8 × 4–6µm
(x̄ = 6.5 × 4.5µm, n = 20), uniseriate, subglobose to ellip-
soidal, aseptate, hyaline when young, becoming pale brown,
smooth-walled, with 2 large guttules, a germ pore at end.
Asexual morph: Undetermined (adapted from Karsten
1873).
Material examined: USA, Louisiana, Ferriday, on dead
wood of Gleditsia sp., 18 July 1931, Overholts and Siggers
(BPI-797104).
Known hosts and distribution: On dead wood of Alnus
glutinosa in UK (type locality) (Karsten 1873); on dead
wood in tropical America and rare in Europe (Miller 1930);
on decayed wood in Argentina (Mercuri 1972).
Notes: Camarops hypoxyloides was introduced by
Karsten (1873). Miller (1930) proposed that C. hypoxyloi-
des is closely related to Solenoplea peltata Lloyd from Porto
Rico, and found that ‘The Montagne specimen of Hypoxylon
polyspermum at Kew is, according to Mason in a letter, the
Fig. 16 Camarops hypoxyloides: a, c–e, g (BPI-797104); b, f (redrawn from Mercuri 1972). a Herbarium material. b, c Ascostromata (b monos-
tichous ascomata). d Ascoma in cross section. e Ostiole. f Ascospores. g Ascus. Scale bars: c = 1mm, d = 100µm, e. f = 50µm, g = 10µm
Fungal Diversity
1 3
same as his English specimen of Camarops which is equal to
Karsten’s C. hypoxyloides’, and then introduced Camarops
polysperma (≡ Hypoxylon polyspermum Mont.) as the cur-
rent name of C. hypoxyloides (Miller 1930; Mercuri 1972).
However, this statement is not generally recognized and
C. hypoxyloides is widely accepted as the type species of
Camarops (Untereiner etal. 2013; Hyde etal. 2020; Index
Fungorum 2020). We also recommend C. hypoxyloides, the
name first used to introduce Camarops, as the type species,
and accept C. polysperma as an independent species similar
to C. hypoxyloides, until there is sufficient morphological
or molecular data to discern their relationship. Raja etal.
(2011) sequenced C. polysperma which was collected from
Sweden. Camarops tubulina is sister to C. polysperma
(95%ML/0.99PP, Fig.12). Morphologically, the former
has polystichous ascomata in stroma, whereas the latter has
monostichous ascomata (Shear 1938; Mercuri 1972). In this
study, we were unable to obtain the type specimen. There-
fore, we re-examined an authentic material (BPI-797104)
collected in America.
Camarops tubulina (Alb. & Schwein.) Shear, Mycologia
30(5): 585 (1938)
Basionym: Sphaeria tubulina Alb. & Schwein., Consp.
fung. (Leipzig): 6 (1805)
Facesoffungi number: FoF 10008; Fig.17
Saprobic on wood. Sexual morph: Ascostromata
15–20 × 2–5 mm (x̄ = 18 × 4 mm, n = 5), carbonaceous
to coriaceous, immersed to semi-immersed, dark brown,
smooth, pulvinate, discoid, hemispherical or irregu-
lar, with broad attachment to the substrate. Ascomata
1–2.5 × 0.5–1mm (x̄ = 1.5 × 0.7mm, n = 10), perithecial,
gregarious, polystichous, carbonaceous, globose to sub-
globose, dark brown to black, ostiolate, with necks. Necks
carbonaceous, dark brown to black, the wall composed of
brown to hyaline cells of textura intricata to textura prismat-
ica, with hyaline periphyses. Peridium of venter 40–120µm
(x̄ = 90µm, n = 30) wide, comprising two layers, outer
layer coriaceous, composed of brown to dark brown cells
of textura angularis; inner layer membranaceous, com-
posed of hyaline cells of textura prismatica. Paraphyses
0.8–1.8µm wide, numerous, septate, branched, filiform. Asci
30–55 × 2–4.5µm (x̄ = 40 × 3µm, n = 50), 8-spored, unitu-
nicate, cylindrical, pedicellate, apex rounded, with J- apical
ring. Ascospores 3–6 × 1.5–2.5µm (x̄ = 4.5 × 2µm, n = 50),
uniseriate, subglobose to ellipsoidal, aseptate, hyaline when
young, becoming dark brown or dark grey, with germ spore
at base, smooth-walled, with 2 guttules. Asexual morph:
Undetermined.
Material examined: Czech Republic, Novohradských
Horách Mountains, Virgin forest of Žofínský prales, on the
dead trunks of Picea sp., 18 October 1967, Z. Pouzar, deter-
mined by M. Svrček in January 1969 (PRM-PR647008).
Known hosts and distribution: On trunk of Abies in Upper
Lusatia (type locality) (Albertini and Schweinitz 1805); on
dead spruce stumps in moist places in the south of Sweden
(Fries 1816); on decayed wood of Abies, rare Juglans and
Alnus in Lusatia (Germany), Sweden and North America
(Saccardo 1882); on dead wood of conifers (Picea, Abies)
and rare broadleaved trees (Fagus) in Czech Republic (Holec
2005).
Notes: Saccardo (1882) transferred Sphaeria tubulina
Alb. & Schw. to Bolinia as generic type (see notes for Bolin-
iaceae). Shear (1938) and Svrček (1969) combined Sphaeria
tubulina Alb. & Schw. under Camarops tubulina (see notes
for Camarops). Thus, Bolinia was subsequently accepted as
a synonym for Camarops, and its members were assigned to
Camaropella and Camarops (Shear 1938; Nannfeldt 1972;
Vasilyeva etal. 2007). Rehm (1904) and Theissen (1909)
proposed C. tubulina is a synonym of C. hypoxyloides, but
Miller (1930) thought that the correct samples were not
examined. Thus, there is an issue with their assumption, that
is, C. tubulina (≡ Sphaeria tubulina) collected by Mason has
polystichous ascomata in stroma with long elongate necks,
different from C. hypoxyloides as described by Karsten has
monostichous ascomata. Huhndorf etal. (2004b) sequenced
C. tubulina, which is sister to C. polysperma (Raja etal.
2011; this study, 95%ML/0.99PP, Fig.12). In this study, we
were unable to obtain the type material. Therefore, we re-
examined an authentic sample determined by Svrček (men-
tioned in the label of material). Camarops hypoxyloides and
C. tubulina are two vital species in Boliniaceae (Boliniales),
and their relationships should be resolved following fresh
collections and epitypification.
Cornipulvina Huhndorf, A.N. Mill., F.A. Fernández &
Lodge 2005
Saprobic on decorticated wood. Sexual morph: Ascos-
tromata superficial, irregular in shape, membranaceous,
brown to black, glabrous, with fascicular arrangement, necks
rostrate. Ascomata perithecial, subglobose, dark brown to
black, ostiolate, with papilla, periphysate. Peridium com-
posed of brown to dark brown cells of textura angularis.
Paraphyses numerous, septate, unbranched, filiform, gut-
tulate, evanescent. Asci 8-spored, unitunicate, cylindrical,
pedicellate, apex rounded, with J- inconspicuous apical ring.
Ascospores uniseriate, subglobose to ellipsoidal, aseptate,
hyaline, smooth-walled, with 2 guttules, without gelatinous
appendages. Asexual morph: Undetermined (adapted from
Huhndorf etal. 2005).
Notes: The monotypic genus Cornipulvina is character-
ized by ascostromata with long rostrate necks and ellipsoi-
dal to subglobose ascospores without germ pores (Huhndorf
etal. 2005). Sequence data from Cornipulvina ellipsoides
(SMH 1378) is basal to Apiorhynchostoma and Pseudovals-
aria in Boliniaceae (Huhndorf etal. 2005; 70%ML, Fig.12).
Fungal Diversity
1 3
Fig. 17 Camarops tubulina – as Bolinia tubulina: (PRM-PR647008).
a Herbarium material. b, c Ascostromata in apical view. d Ascos-
troma in longitudinal section. e Ascomata in cross section. f Neck
vertical section with periphyses. g Peridium of the base. h, i Para-
physes (h mounted in cotton blue). j–m Asci. n–q Apical ring
of ascus. r–u Ascospores (s arrow points to germ pore). Scale
bars: b. e = 500 µm, c = 200 µm, d = 2 mm, f = 50 µm, g = 20 µm,
h–m = 10µm, n–q = 5µm, r–u = 2µm
Type species: Cornipulvina ellipsoides Huhndorf, A.N.
Mill., F.A. Fernández & Lodge, Fungal Diversity 20: 63
(2005)
Facesoffungi number: FoF 10009; Fig.18
Saprobic on wood. Sexual morph: Ascostromata
1.5–3.5 × 1–2.5 mm (x̄ = 2.5 × 1.5 mm, n = 5), superfi-
cial, irregular in shape, membranaceous, brown to black,
glabrous, with fascicular arrangement, necks rostrate. Asco-
mata 450–900 × 250–600 µm (x̄ = 750 × 450µm, n = 10),
perithecial, subglobose, dark brown to black, ostiolate,
with papilla, periphysate. Peridium 25–65µm (x̄ = 50µm,
n = 30) wide, composed of brown to dark brown cells of
textura angularis. Paraphyses 1–3.5µm wide, numerous,
septate, unbranched, filiform, guttulate, evanescent. Asci
Fungal Diversity
1 3
(50–)60–77(–85) × 3.5–7 µm (x̄ = 70 × 5.5 µm, n = 50),
8-spored, unitunicate, cylindrical, pedicellate, apex
rounded, with J- inconspicuous apical ring. Ascospores
(5–)7–8.5(–10) × 3–5µm (x̄ = 8 × 4.5µm, n = 50), uniseriate,
subglobose to ellipsoidal, aseptate, hyaline, smooth-walled,
usually with two large guttules, without gelatinous append-
ages. Asexual morph: Undetermined.
Material examined: Puerto Rico, Luquillo Mountains,
Bisley Watershed 3, 220m, 18º 19.00′ N/65° 0′ W, on dead
decorticated wood, 8 May 1995, S.M. Huhndorf and D.J.
Lodge (F-SMH 1378, holotype).
Known hosts and distribution: On decayed decorticated
wood in Brazil, Puerto Rico (type locality) and Venezuela
(Huhndorf etal. 2005).
Notes: We re-examined the holotype of Cornipulvina
ellipsoides (SMH 1378), and it is sister to Apiorhynchostoma
and Pseudovalsaria (70%ML, Fig.12).
Endoxyla Fuckel, Jb. nassau. Ver. Naturk. 25–26: 321 (1871)
Saprobic on wood. Sexual morph: Ascostromata poorly
developed or absent. Ascomata perithecial, gregarious or in
bi-seriate linear groups, immersed to semi-immersed, glo-
bose to subglobose, black, ostiolate. Asci 8-spored, clavate,
pedicellate. Ascospores ellipsoidal to cylindrical, hyaline
to brown, slightly curvate, with a germ pore at one end.
Asexual morph: Undetermined (adapted from Fuckel 1872).
Notes: Fuckel (1872) established Endoxyla based on glo-
bose ascomata with clavate asci and ellipsoidal ascospores,
and while introducing E. macrostoma, mentioned that Spha-
eria parallela Fr. and S. operculata Alb. & Schw. belong to
Endoxyla. Saccardo (1882) accepted S. parallela as Endox-
yla parallela and S. operculata as E. operculata. Subse-
quently, there was a controversy as to which species should
be selected as the type of Endoxyla. Some mycologists have
recognized S. operculata (≡ E. operculata) published earlier,
Fig. 18 Cornipulvina
ellipsoides: (F-SMH 1378,
holotype). a Herbarium
material label. b, c Cluster of
ascomata on host. d Ascoma
in cross section. e, f Asci
with paraphyses. g–i Apical
ring. j–n Ascospores. Scale
bars: c = 1mm, d = 200µm, e.
f = 50µm, g–n = 5µm
Fungal Diversity
1 3
as the type species (Clements and Shear 1931; Holm 1975).
Others believed that Fuckel (1872) only described E. mac-
rostoma in detail when establishing Endoxyla, so that, it
should be regarded as the generic type (Untereiner 1993).
Endoxyla macrostoma is widely accepted as the type spe-
cies of Endoxyla (Untereiner 1993; Untereiner etal. 2013;
Hyde etal. 2020). Untereiner etal. (2013) sequenced strains
of E. macrostoma, E. mallochii and E. operculata which
clustered with Endoxyla (98%ML, Fig.12) in Boliniaceae.
The Endoxyla species forms a sister clade to Camaropella
(53%ML, Fig.12).
Type species: Endoxyla macrostoma Fuckel, Jb. nassau.
Ver. Naturk. 25–26: 322 (1871)
Facesoffungi number: FoF 10119; Fig.20l–m.
Saprobic on decorticated wood. Sexual morph: Ascos-
tromata poorly developed, brown, septate hyphae. Ascomata
perithecial, gregarious or in bi- or uni-seriate linear groups,
immersed to semi-immersed, ampulliform to subglobose,
black, ostiolate, with papilla. Peridium comprising two lay-
ers, outer layer composed of dark brown to brown cells of
textura intricata to textura angularis; inner layer composed
of hyaline cells of textura prismatica. Paraphyses filiform,
septate, sparse in mature ascomata. Asci 8-spored, clavate
to cylindrical, pedicellate, with J- apical ring. Ascospores
ellipsoidal to suballantoid, curved, 0–1-septate, unequal,
composed of a large brown to pale brown cell with an apical
germ pore and a tiny, conical, hyaline cell. Asexual morph:
Undetermined (adapted from Fuckel 1872; Untereiner 1993).
Known hosts and distribution: On decayed, decorticated
wood of Quercus in Germany (type locality) (Fuckel 1872);
on decorticated wood of angiosperms and gymnosperms
(Abies, Picea, Pinus) in Canada, Czech Republic, Germany
and Sweden (Untereiner 1993; Untereiner etal. 2013).
Notes: Fuckel (1872) established Endoxyla and introduced
E. macrostoma which was collected from Germany. Holm
(1975) suggested E. operculata as the type species. However,
Untereiner (1993), after reviewing the species of Endoxyla,
proposed that E. macrostoma should be used as the type (see
notes for Endoxyla). Untereiner etal. (2013) sequenced E.
macrostoma collected from Canada and, this strain is basal
to E. mallochii and E. operculata (84%ML, Fig.12).
Endoxyla operculata (Alb. & Schw.) Sacc., Syll. fung.
(Abellini) 1: 181 (1882)
Basionym: Sphaeria operculata Alb. & Schwein., Consp.
fung. (Leipzig): 40 (1805)
Facesoffungi number: FoF 10010; Fig.19
Saprobic on wood. Sexual morph: Mycelium brown,
septate hyphae 2–6.5µm wide, surrounded the base of the
ascoma. Ascostromata poorly developed or absent. Asco-
mata 800–900 × 500–565 µm (x̄ = 845 × 530µm, n = 10),
perithecial, solitary, immersed, with necks erumpent the
rough wood of the surface, in linear groups, ampulliform to
subglobose, black, ostiolate, with necks, periphysate. Perid-
ium 40–100µm (x̄ = 70µm, n = 30) wide, comprising three
layers, outer layer carbonaceous or coriaceous, composed
of dark brown cells of textura intricata; middle layer mem-
branaceous, composed of brown cells of textura angularis;
inner layer membranaceous, composed of hyaline cells of
textura prismatica. Paraphyses filiform to ventricose, sep-
tate, sparse in mature ascomata. Asci 85–90 × 6.5–12 µm
(x̄ = 87.5 × 9.5µm, n = 30), 8-spored, unitunicate, cylindri-
cal to clavate, pedicellate, apex blunt, with J- apical ring.
Ascospores 9.5–16 × 4–6.5 µm (x̄ = 13.5 × 5µm, n = 50),
uni- or bi-seriate, ellipsoidal to cylindrical, aseptate, hyaline
when young, becoming pale grey, smooth-walled, slightly
curved, guttulate, sometimes with an apical germ pore.
Asexual morph: Undetermined (adapted from Untereiner
1993).
Material examined: Sweden, Uppland, Östera Ryd par.,
Rydboholm. S. of the castle in deciduous forest, on naked
wood of deciduous tree, 17 May 1983, N. Lundqvist (S-F
138814); Norway, Troms, Storord commune, Helligsko-
gen in Skibotndalen Valley (26km SE of Skibotn), subal-
pine birch forest, on decayed wood of Betula pubescens, 17
August 1992, N. Lundqvist (S-F 138817).
Known hosts and distribution: On decayed wood of
Abies in Lusatia and Sweden (type locality) (Albertini and
Schweinitz 1805); on decayed, decorticated wood of angio-
sperms (Acer, Fagus, Populus, Quercus) and gymnosperms
(Alnus, Pinus) in Canada, Czech Republic, Finland, France,
Germany, Sweden, USA (Untereiner 1993; Untereiner etal.
2013).
Notes: Untereiner (1993) introduced a lectotype for
Endoxyla operculata which was collected from Sweden by
Fries which has immersed ascomata with clavate to cylin-
drical asci and cylindrical ascospores; however, the stomata
are poorly developed or absent. In this study, we could not
obtain the type material. Therefore, we re-examined authen-
tic specimens collected by Lundqvist in Sweden and Nor-
way. These collections of E. operculata examined here, had
prominent apical rings in the asci, and ascomata were not
in a stroma (Fig.19). These characters are not identical to
Camarops tubulina, which is considered to be the current
name of E. operculata (Index Fungorum 2020). Endoxyla is
different from Camarops with crust-like stroma. Phylogenet-
ically, the strain of E. operculata is sister to E. mallochii and
is distant from C. tubulina (Untereiner etal. 2013; Réblová
etal. 2018; this study, 84%ML, Fig.12).
Mollicamarops Lar.N. Vassiljeva, Mycotaxon 99: 160
(2007)
Saprobic on decorticated wood. Sexual morph: Ascostro-
mata semi-immersed, irregular in shape, widely effuse, ochre-
ous or fulvous. Ascomata black, ostiolar region conspicuously
Fungal Diversity
1 3
stellate to furrowed. Paraphyses numerous, filiform, sinuous.
Asci 8-spored, unitunicate, cylindrical, apical ring indistinct,
long pedicellate. Ascospores ellipsoidal, light brown to brown,
aseptate, smooth-walled, with guttules. Asexual morph:
Undetermined (adapted from Vasilyeva 2007).
Notes: The monotypic genus Mollicamarops is charac-
terized by stellate, ostiolate ascomata, cylindrical asci and
ellipsoidal, aseptate ascospores. It is similar to Camarops
flava in having yellow to brown ascostromata and can be
distinguished as the latter has astellate ostioles and orna-
mented ascospores (Samuels and Rogers 1987; Vasilyeva
2007). Therefore, we place Mollicamarops in Boliniaceae.
Type species: Mollicamarops stellata Lar.N. Vassiljeva,
Mycotaxon 99: 160 (2007)
Facesoffungi number: FoF 10120; Fig.20 n–o
Saprobic on decorticated wood. Sexual morph: Ascos-
tromata semi-immersed, irregular in shape, widely effuse,
ochreous or fulvous. Ascomata black, ostiolar region con-
spicuously stellate to furrowed. Paraphyses numerous, nar-
row, sinuous. Asci 8-spored, unitunicate, cylindrical, apical
ring indistinct. Ascospores ellipsoidal, light brown, aseptate,
smooth-walled, with guttules. Asexual morph: Undeter-
mined (adapted from Vasilyeva 2007).
Known hosts and distribution: On decayed, decorticated
wood in Russia (type locality) (Vasilyeva 2007).
Notes: We were unable to obtain type material. Therefore,
we draw the asci and ascospores of Mollicamarops stellata
(Fig.20n–o) from Vasilyeva (2007).
Neohypodiscus J.D. Rogers, Y.M. Ju & Læssøe, Mycologia
86(5): 684 (1994)
Fig. 19 Endoxyla operculata:
a, b, e–f, j, l, s (S-F 138,817);
c, d, g–i, k, m–r (S-F 138,814).
a Herbarium material. b,
c Ascomata on host. d, e
Ascomata in cross section. f,
g Peridium. h Neck in cross
section with periphyses. i Base
of ascoma with mycelium. j–m
Asci. n–s Ascospores. Scale
bars: b–c = 1mm, d. i = 200µm,
e = 500µm, f. g = 100µm, h.
j–m = 50µm, n–s = 10µm
Fungal Diversity
1 3
Saprobic on decorticated wood. Sexual morph: Ascostro-
mata subglobose to pulvinate to irregular, solitary or conflu-
ent, narrow to broad, carbonaceous, with longitudinal veins
below and attachment to the substrate. Ascomata subglobose
to ampulliform, brown to black, subglobose, carbonaceous,
monostichous, ostiolate, rough-walled. Paraphyses numer-
ous, mostly unbranched, septate. Asci 8-spored, unitunicate,
cylindrical to clavate, with J-apical ring. Ascospores uni-
seriate, greenish to brown, ellipsoidal to crescent-shaped,
0–2-septate, smooth-walled, some with an apical thickening
or acellular appendage at one end, with a germ pore at one
or both end(s), guttulate. Asexual morph: Undetermined
(adapted from Rogers etal. 1994b).
Notes: Rogers etal. (1994b) placed Neohypodiscus in
Amphisphaeriaceae (Xylariomycetidae) based on its carbo-
naceous stromata, cylindrical asci and ellipsoidal to fusiform
ascospores. Subsequently, Andersson etal. (1995) trans-
ferred it to Boliniaceae. No molecular data for this genus
Fig. 20 Neohypodiscus rickii:
a, c, e, g, h (BPI-737826);
b, d, f, i (BPI-802543); j, k
(redrawn from Rogers etal.
1994b). a, b Herbarium mate-
rial. c, d Ascostromata on host.
e, f Ascostromata narrow to
broad connection to substrate
with veins. g–i Ascomata (h
arrow indicates short umbili-
cate ostiole). j Ascospores
with appendage at the ends. k
Ascus with indistinct apical
ring; Endoxyla macrostoma:
l, m (redrawn from Untereiner
1993). l Ascus. m Ascospores;
Mollicamarops stellata: n,
o (redrawn from Vasilyeva
2007). n Ascus. o Ascospores;
Pseudovalsaria foedans: p–q
(redrawn from Spooner 1986).
p Ascus. q Ascospores. Scale
bars: c–f = 2mm, g–i = 1mm,
j. m–n. q = 10µm, k = 50µm, l,
p = 20µm, o = 5µm
Fungal Diversity
1 3
has been sequenced yet. In this study, we maintain Neohypo-
discus in Boliniaceae because it has the same carbonaceous
stromata and unequal ascospores with hyaline thickened
end(s) as Camarops and Apiorhynchostoma.
Type species: Neohypodiscus rickii (Lloyd) J.D. Rogers,
Y.M. Ju & Læssøe, Mycologia 86(5): 685 (1994)
Basionym: Hypodiscus rickii Lloyd, Mycol. Writ. 7(Let-
ter 68): 1181 (1923)
Facesoffungi number: FoF 10,121; Fig.20a–k
Saprobic on decorticated wood. Sexual morph: Ascostro-
mata subglobose to irregular, confluent in mass, with narrow
to broad connection to substrate, brown to black, with veins.
Ascomata 350–1300 × 550–800 µm (x̄ = 1100 × 755 µm,
n = 10), brown to black, subglobose, carbonaceous, with
a short umbilicate ostiole, rough-walled. Paraphyses
1.5–10µm wide, numerous, mostly unbranched, septate.
Asci 200 × 11µm, 8-spored, unitunicate, cylindrical, api-
cal ring indistinct. Ascospores 26.5–32 × 7–9µm, uniseri-
ate, greenish to brown, ellipsoidal to crescentic, aseptate or
1-septate near one end or rarely 2-septate, unequal, normally
composed of one small cell and one large cell with a germ
pore at end, smooth-walled, some with an apical thickening
or acellular appendage at one end, with guttules. Asexual
morph: Undetermined (adapted from Rogers etal. 1994b).
Material examined: Puerto Rico, El Toro Trail, off Rt.
186, Luquillo Mountains, on dead wood, 24 December 1996,
D.J. Lodge, G.J. Samuels and H.J. Strauss (BPI-737826);
Puerto Rico, El Verde, Luquillo Mts., Trail to the Radiation
Center, on dead wood, 23 March 1993, D.J. Lodge and M.
Boyd (BPI-802543).
Known hosts and distribution: On decayed, decorticated
wood in Brazil (type locality), Puerto Rico (Rogers etal.
1994b).
Notes: Lloyd (1923) described Hypodiscus rickii, and
Rogers etal. (1994b) transferred this species as Neohypo-
discus rickii as its generic type. Rogers etal. (1994b) exam-
ined two samples collected by Lloyd from Brazil and defined
those collections as lectotype and paralectotype materials.
In this study, we re-examined two authentic samples deter-
mined by Rogers (mentioned in the label of material) as we
were unable to obtain type material. We were unable to find
any structures in the samples except the ascomata, there-
fore, we provided a hand drawing of asci and ascospores
(Fig.20j–k) following Rogers etal. (1994b).
Pseudovalsaria Spooner, Trans. Br. mycol. Soc. 86(3): 405
(1986)
Saprobic on wood. Sexual morph: Ascostromata
immersed to erumpent through bark. Ascomata gregarious,
brown to black, ellipsoidal to pyriform, immersed to semi-
immersed in stroma. Asci 8-spored, unitunicate, cylindrical,
apex rounded or blunt. Ascospores uniseriate, brown to dark
brown, cylindrical to ellipsoidal, ends slightly tapered to
rounded, 0–1-septate, smooth-walled, with a germ pore at
the base, guttulate. Asexual morph: Undetermined (adapted
from Spooner 1986).
Notes: Spooner (1986) established Pseudovalsaria typi-
fied by P. foedans, and Barr (1994) placed this genus in
Clypeosphaeriaceae. Rappaz (1995) subsequently trans-
ferred it to Boliniaceae as it has similar characters to Cama-
rops, such as cylindrical asci and ellipsoidal ascospores
with germ pore. Untereiner etal. (2013) confirmed that
Pseudovalsaria belong in Boliniaceae based on ITS-LSU
sequence analyses of P. ferruginea. In this study, the strain
of P. ferruginea is sister to Apiorhynchostoma apiosporum
in Boliniaceae (72%/ML, Fig.12).
Type species: Pseudovalsaria foedans (P. Karst.)
Spooner, Trans. Br. mycol. Soc. 86(3): 405 (1986)
Basionym: Phaeosperma foedans P. Karst., Bidr. Känn.
Finl. Nat. Folk 23: 55 (1873)
Facesoffungi number: FoF 10122; Fig.20p–q
Saprobic on wood of Pinus silvestris. Sexual morph:
Ascostromata gregarious, immersed to erumpent through
bark. Ascomata gregarious, immersed to semi-immersed in
stroma, brown to black, ellipsoidal to pyriform, with long
necks. Necks composed of pale brown to hyaline cells of
textura porrecta, surrounded by stromatic tissue. Peridium
of venter outer layer composed of dark brown, irregular
patches forming opaque amorphous matter; middle layer
composed of brown cells of textura angularis to textura
prismatica; inner layer composed of hyaline cells of textura
prismatica. Paraphyses mostly branched, filiform, septate.
Asci 8-spored, unitunicate, cylindrical, apex rounded, with
J- apical ring. Ascospores uniseriate, dark brown, cylindrical
to ellipsoidal, slightly tapered to rounded ends, sometimes
slightly curved, 1-septate, smooth-walled, with a termi-
nal germ pore, guttulate. Asexual morph: Undetermined
(adapted from Spooner 1986).
Known hosts and distribution: On decorticated wood of
Viburnum sp. in China (Spooner 1986); on decayed wood
of Betula sp., Castanea sativa, Pinus silvestris, Prunus lau-
rocerasus, Quercus sp. in England (type locality) (Spooner
1986); on decayed wood of Alnus in Finland (Spooner
1986); on decayed wood of Alnus sp. in Canada (Untereiner
etal. 2013).
Notes: Spooner (1986) established Pseudovalsaria
based on Phaeosperma foedans. Untereiner (1993) pro-
posed that Anthostoma ferrugineum, which has J-asci and
brown, 1-septate ascospores with an apical germ pore, is
similar to Pse. foedens. Rappaz (1995) subsequently syn-
onymized A. ferrugineum under Pse. ferruginea and men-
tioned that ascospores of Pse. foedans are normally cylin-
drical and slightly curved which distinguishes it from Pse.
ferruginea which has symmetrical, cylindrical ascospores.
Ju etal. (1996) believed that the difference in ascospores
between P. ferruginea and P. foedans were indistinct, and
Fungal Diversity
1 3
these species were probably conspecific. They also merged
these two species and recognized P. ferruginea as the cur-
rent name (Ju etal. 1996). However, this was not verified
by phylogenetic analysis. Morphologically, Rappaz (1995)
had considered Pse. ferrugine as having grey-brown tissue
around the brown–red ascomata as different to Pse. foedans,
which has dark brown tissue around the dark brown asco-
mata (Spooner 1986). The holotype of Pse. ferruginea and
Pse. foedans were collected from Germany and England
respectively (Nitschke 1867; Karsten 1873). Untereiner etal.
(2013) analyzed the sequence data from a collection of Pse.
ferruginea which was isolated from a decorticate branch of
Populus in Canada. This study recommends that the generic
type Pse. foedans (Spooner 1986) and Pse. ferruginea as two
independent species until there are more characteristics and
phylogenetic data to determine their classification. The strain
of Pse. ferruginea is sister to Apiorhynchostoma apiosporum
(72%/ML, Fig.12), but no sequence is available for Pse.
foedans. We were unable to examine the type taxon, there-
fore, we provided a hand drawing of asci and ascospores
(Fig.20p–q) following Spooner (1986).
Chaetosphaeriales Huhndorf, A.N. Mill. & F.A. Fernández,
Mycologia 96(2): 378 (2004)
Chaetosphaeriales was introduced by Huhndorf etal.
(2004a) and four families (Chaetosphaeriaceae, Hel-
minthosphaeriaceae, Leptosporellaceae and Linocarpaceae)
are included in this order based on the morphology and
phylogenetic analyses (Maharachchikumbura etal. 2016;
Hernández-Restrepo etal. 2017; Konta etal. 2017; Hyde
etal. 2020). Species of Chaetosphaeriales are mostly sap-
robes or/and pathogens in terrestrial and aquatic habitats,
and can also be fungicolous (Maharachchikumbura etal.
2015, 2016).
Helminthosphaeriaceae Samuels, Cand. & Magni, Myco-
logia 89(1): 144 (1997)
Saprobic or parasitic on wood or decorticated branches
or stems in terrestrial habitats, sometimes fungicolous.
Sexual morph: Ascomata perithecial, solitary, scattered or
gregarious, superficial or immersed, ovoid to subglobose,
dark brown to black, carbonaceous or membranaceous,
glabrous or tuberculate or with setae, papillate. Peridium
carbonaceous or membranaceous, composed of brown to
hyaline cells of textura angularis to textura prismatica.
Paraphyses numerous, evanescent, septate, unbranched or
branched, filiform to cylindrical. Asci 8-spored, unituni-
cate, cylindrical to clavate, apex truncate, pedicellate, with
J-apical ring. Ascospores normally bi-seriate, allantoid or
clavate or cylindrical, 0–3-septate, hyaline to brown or grey
or becoming dark colored in part, smooth-walled, with germ
pore at one end or/and both ends, with guttules or guttules
absent. Asexual morph: Hyphomycetous. Conidiophores
macronematous, mononematous, gregarious, cylindrical,
branched or unbranched, pale brown to brown, septate,
smooth-walled. Conidia solitary, acrogenous, ellipsoidal to
obclavate, aseptate to multi-septate, smooth-walled, hyaline
to brown, with guttules or guttules absent (adapted from
Fuckel 1870; Réblová 1999b; Maharachchikumbura etal.
2016).
Type genus: Helminthosphaeria Fuckel 1870
Notes: Helminthosphaeriaceae is typified by Hel-
minthosphaeria (Fuckel 1870) and most of its species have
black setose, globose ascomata and cylindrical asci with
brown to hyaline, subglobose ascospores and mononema-
tous conidiophores with oblong conidia. Initially, Echino-
sphaeria, Endophragmiella, Helminthosphaeria, Hilberina,
Ruzenia and Synaptospora were included based on morpho-
logical and/or phylogenetic analysis (Miller and Huhndorf
2004a; Miller etal. 2014; Maharachchikumbura etal. 2015).
Diplococcium and Tengiomyces were added to this family
(Lumbsch and Huhndorf 2007; Wijayawardene etal. 2018).
Tengiomyces was accepted in this family as its ascomata
are similar to Helminthosphaeria (Réblová 1999a), whereas
its ascospores are similar to Chaetosphaerella and Cras-
sochaeta (Chaetosphaerellaceae, Coronophorales) (Réblová
1999a; Hyde etal. 2020; Wijayawardene etal. 2020). Thus,
we suggest placing Tengiomyces in Coronophorales genera
incertae sedis (see notes for Tengiomyces). The hyphomy-
cetous Diplococcium and Endophragmiella asexual morphs
have associated species of Helminthosphaeria and Echino-
sphaeria on the natural substrate (Sutton 1973; Samuels
etal. 1997; Goh and Hyde 1998; Goh etal. 1998; Réblová
1999b; Huhndorf etal. 2004b). Diplococcium has phylo-
genetic affinities with Helotiales (Leotiomycetes) and Ple-
osporales (Dothideomycetes) (Shenoy etal. 2007, 2010);
and Endophragmiella species are closely related to Hel-
minthosphaeriaceae and Tubeufiales (Dothideomycetes)
(Hernández-Restrepo etal. 2017; Vu etal. 2019). However,
Diplococcium was transferred to Vibrisseaceae (Ekanayaka
etal. 2019) and we recommend to place Endophragmiella
in Ascomycota genera incertae sedis (see notes for Endo-
phragmiella). In this study, Synaptospora is transferred to
Sordariales genera incertae sedis (see notes for Synaptos-
pora). Thus, four genera, Echinosphaeria, Helminthospha-
eria, Hilberina and Ruzenia are presently accepted in
Helminthosphaeriaceae.
Echinosphaeria A.N. Mill. & Huhndorf, Mycol. Res.
108(1): 29 (2004)
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, solitary or scattered, superficial, subglobose to ovoid,
membranaceous, brown to black, with setae, ostiolate, with
papilla, periphysate. Setae hyaline to brown, straight, rigid,
with acute apex. Peridium composed of loose arrangement,
of hyaline to brown cells of textura angularis to textura
Fungal Diversity
1 3
prismatica. Asci 8-spored, unitunicate, cylindrical to clavate,
apex blunt, with J-apical ring, pedicellate. Ascospores cylin-
drical to allantoid, aseptate to multi-septate, hyaline to brown,
smooth-walled or verrucose, ends rounded, slightly curved,
guttulate. Asexual morph: Undetermined (adapted from
Persoon 1796; Karsten 1873; Miller and Huhndorf 2004a).
Notes: Miller and Huhndorf (2004a) established
Echinosphaeria for Lasiosphaeria canescens based on
LSU sequence analysis. This genus clustered with Hel-
minthosphaeria in Helminthosphaeriaceae (Hongsanan
etal. 2017; Wijayawardene etal. 2018; Hyde etal. 2020). In
this study, Echinosphaeria nests in Helminthosphaeriaceae
based on the LSU-TUB sequence analysis (Fig.21).
Type species: Echinosphaeria canescens (Pers.) A.N.
Mill. & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Basionym: Sphaeria canescens Pers., Observ. mycol.
(Lipsiae) 1: 67 (1796)
Facesoffungi number: FoF 10011; Fig.22a–r
Saprobic on wood. Sexual morph: Ascomata
365–660 × 340–590µm (x̄ = 470 × 450µm, n = 10), per i-
thecial, solitary or scattered, superficial, globose to ovoid,
membranaceous, brown to black, with brown, densely
setae, ostiolate, with papilla, periphysate. Setae 5.5–12µm
wide at base, rigid, hyaline to pale brown, with acute apex.
Peridium 25–50µm (x̄ = 37µm, n = 30) wide, comprising
two layers, outer layer composed of loose arrangement,
of brown to pale brown cells of textura angularis; inner
layer composed of pale brown to hyaline cells of textura
prismatica. Paraphyses 2–4µm wide, numerous, septate,
unbranched, filiform. Asci 105–115(–125) × 9–15.5 µm
0.03
Endophragmiella dimorphospora FMR 12150
Hilberina munkii SMH1531
Coniochaeta discoidea SANK12878
Helminthosphaeria ludens JF04126
Helminthosphaeria odontiae ANM928
Helminthosphaeria triseptata JF04015
Synaptospora plumbea ANM963
Coniochaeta ligniaria SMH2569
Helminthosphaeria clavariarum ANM Acc.17
Helminthosphaeria carpathica SMH3903
Helminthosphaeria corticiorum JF04225
Helminthosphaeria tomaculum SMH2485(F)
Ruzenia spermoides SMH4655
Ruzenia spermoides SMH4606
Hilberina robusta SMH3054
Hilberina sphagnorum Buck49156
Ruzenia spermoides CBS 101621
Echinosphaeria canescens SMH4666
Echinosphaeria canescens JHC97-006
Helminthosphaeria hyphodermae SMH4192
Ruzenia spermoides ANM163
Hilberina punctata SMH4825
Helminthosphaeria mammillata ANM986
Hilberina caudata SMH1542
Synaptospora plumbea SMH3962
Echinosphaeria canescens TL5730
Coniochaetidium savoryi TRTC51980
61/--
95/0.95
96/1.00
85/0.95
57/--
100/1.00
90/0.93
100/1.00
88/0.99
93/0.92
100/1.00
100/1.00
100/1.00
Coniochaetales
Helminthosphaeriaceae
54/--
54/--
55/--
69/0.93
83/--
Hel. plumbea
Fig. 21 Phylogram generated from maximum likelihood analysis
based on combined LSU and TUB sequence data of Helminthospha-
eriaceae. 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 correspond-
ing nodes. The ‘--’ indicates lack of statistical support (< 50% for
ML-BS and < 0.90 for BY-PP). Twenty-seven strains are included in
the combined analyses which comprise 1648 characters (995 char-
acters for LSU, 653 characters for TUB) after alignment. Strains of
Coniochaetales are used as the outgroup taxa. The best score in IQ-
TREE explores with a final likelihood value of -7018.1436 is pre-
sented. The model of each partitioned gene is: LSU: GTR + I + G;
TUB: GTR + G. The strain numbers are noted after the species
names. Ex-type strains are in bold. Alignments are available at Tree-
BASE (URL: http:// purl. org/ phylo/ treeb ase/ phylo ws/ study/ TB2:
S28267)
Fungal Diversity
1 3
(x̄ = 110 × 12.5µm, n = 30), 8-spored, unitunicate, cylindri-
cal to clavate, pedicellate, apex blunt, with J- apical ring.
Ascospores (22.5–)24–28(–31) × 4–7.5µm (x̄ = 26 × 6µm,
n = 50), bi-seriate, subglobose to allantoid, 0–1-septate, hya-
line to pale brown, smooth-walled, ends rounded, slightly
curved, guttulate. Asexual morph: Undetermined.
Material examined: USA, Michigan, Marquette, Huron
Mountain Club, between Conway Lake and Lake Superior
on road to Todd property, 46º 00′N/87º 0′ W, on driftwood
half buried in sand, 15 July 1996, S.M. Huhndorf etal.
(F-SMH 2627). Sweden, Uppsala, Uppland, on dead wood,
26 August 1884, L. Romell (S-F 134887). Finland, on dead
wood, 16 October 1861, P.K. Karsten (S-F 134901).
Known hosts and distribution: On dead, decorticated
wood of Betula, Pinus and Quercus in Finland (type locality)
(Persoon 1796; Karsten 1873); on dead wood in Denmark,
Sweden and USA (Miller and Huhndorf 2004a).
Notes: Echinosphaeria canescens is the type species of
Echinosphaeria (Miller and Huhndorf 2004a). This species
has been reported with hyphomycetous Endophragmiella
and selenosporella-like. Cultural characteristics for Ech.
canescens are lacking (Hughes 1979; Sivanesan 1983; Miller
Fig. 22 Echinosphaeria canescens: a, b, e, h, i, l–n (F-SMH 2627);
f, g, j, o, p (S-F 134887); c, d, k, q, r (S-F 134901). a Ascomata on
host. b–d Solitary or scattered ascomata. e, f Ascomata in cross sec-
tion with setae. g Peridium. h Seta. i Asci with paraphyses. j–l Asci.
m–r Ascospores; Endophragmiella pallescens: s (redrawn from Sut-
ton 1973). s Conidiophores with conidia. Notes: i. k. Arrows points
to the apical ring of ascus. Scale bars: b–d = 500µm, e. f = 200µm,
g = 100µm, h. m–s = 20µm, i–l = 50µm
Fungal Diversity
1 3
and Huhndorf 2004a; Hernández-Restrepo etal. 2017; Vu
etal. 2019). Strains of Ech. canescens clustered with Hel-
minthosphaeria species in Helminthosphaeriaceae (Miller
etal. 2014; Miller and Huhndorf 2004a; Hongsanan etal.
2017; this study, Fig.21). The molecular data of two Endo-
phragmiella species are available, End. taxi is closely related
to Tubeufiales (Dothideomycetes) (Vu etal. 2019); and End.
dimorphospora is nested in Helminthosphaeriaceae, but is
phylogenetically distant from Ech. canescens (Hernández-
Restrepo etal. 2017; this study, Fig.21). Selenosporella
strains are more closely related to Diatrypaceae (Xylariales)
(Vu etal. 2019). In this study, we are unable to obtain the
type material. We re-examined authentic samples collected
by Karsten and Hundorf.
Helminthosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23-24:
166 (1870) [1869-70]
Saprobic or parasitic on wood or decorticated branch in
terrestrial habitats, sometimes fungicolous. Sexual morph:
Ascomata perithecial, solitary, scattered or gregarious,
superficial or immersed, globose to subglobose, dark brown
to black, carbonaceous or membranaceous, glabrous or with
setae, with or without ostiole, with periphyses. Peridium
carbonaceous or membranaceous, composed of brown to
hyaline cells of textura angularis to textura prismatica.
Paraphyses numerous, evanescent, septate, unbranched or
branched, filiform to cylindrical. Asci 8-spored, unituni-
cate, cylindrical to clavate, pedicellate, apex truncate, with
J- apical ring. Ascospores bi-seriate, ellipsoidal to clavate,
0–3-septate, hyaline to brown or grey, smooth-walled, with
germ pore at one end or/and both ends, guttulate. Asexual
morph (associated): Hyphomycetous. Conidiophores
macronematous, mononematous, gregarious, cylindrical,
branched, pale brown to brown, septate, smooth-walled.
Conidia solitary, ellipsoidal to obclavate, aseptate to multi-
septate, smooth-walled, hyaline to brown, guttulate (adapted
from Fuckel 1870; Samuels etal. 1997; Miller etal. 2014).
Notes: Desmazières (1837) introduced a species growing
on Clavaria fuliginea in northern France, and Fuckel (1870)
later accepted it as the generic type of Helminthosphaeria,
named as H. clavariarum. This genus was considered as
member of Sordariales based on the ontogeny of the asco-
mata and 1-septate ascospores with germ pores are similar
to Sordaria (Parguey-Leduc 1960; Lundqvist 1972; Eriks-
son and Hawksworth 1993). Subsequently, Samuels etal.
(1997) monographed this genus, established Helminthospha-
eriaceae and concluded that most species in this family
were found on members of Clavariaceae or/and coriaceous
basidiomycetes. They also found that Helminthosphaeria
species were accompanied with hyphomycetous Diplo-
coccium or/and heteroconium-like asexual morphs on the
substrate (Samuels etal. 1997). Afterward, Miller etal.
(2014) introduced several lignicolous Helminthosphaeria
species based on phylogenetic analysis and reported that
the polyphyletic Helminthosphaeria has Diplococcium
and Selenosporella asexual morphs in nature, but not con-
firmed by cultural characteristics or phylogenetic analysis.
Helminthosphaeria is characterized by a fungicolous and
lignicolous habit, superficial or immersed ascomata, cylin-
drical asci with hyaline to brown ascospores and associated
asexual morphs with smooth-walled or verrucose, septate
conidia (Samuels etal. 1997; Goh and Hyde 1998; Réblová
1999a; Hernández-Restrepo etal. 2017). In this study, the
polyphyletic Helminthosphaeria is related to Endophrag-
miella dimorphospora and Hilberina species, and scattered
in Helminthosphaeriaceae (Fig.21). However, we could not
confirm the asexual morph of Helminthosphaeria due to lack
of cultural characteristics and molecular data.
Type species: Helminthosphaeria clavariarum (Desm.)
Fuckel, Jb. nassau. Ver. Naturk. 23-24: 166 (1870) [1869-70]
Basionym: Peziza clavariarum Desm., Annls Sci. Nat.,
Bot., sér. 2 8: 8 (1837)
Facesoffungi number: FoF 10012; Fig.23
Saprobic or parasitic on Clavulina. Sexual morph: Asco-
mata 295–390 × 195–275 µm (x̄ = 335 × 240µm, n = 10),
perithecial, solitary, superficial, globose, black, rough,
surrounded by setae, ostiolate, with papilla, periphysate.
Setae 9.5–15µm wide. brown to pale brown, tapering, with
a septate lumen. Ostioles inconspicuous, a loose arrange-
ment of cylindrical to oblong cells. Peridium 25–50µm
wide, membranaceous, comprising two layers, outer layer
composed of brown cells of textura angularis; inner layer
composed of hyaline cells of textura prismatica. Paraphy-
ses 3–7µm wide, numerous, evanescent, septate, filiform.
Ascogenous hyphae discrete, smooth-walled, branched, with
hyaline, ovoid to obpyriform cells, 9.5–12 × 3.5–6.5µm
(x̄ = 11 × 5.5 µm, n = 10), growing in sympodial suc-
cession, in dense clusters, each giving rise to an ascus.
Asci (70–)80–85(–110) × 8–13.5 µm (x̄ = 82 × 10 µm,
n = 30), 8-spored, unitunicate, cylindrical, apex rounded,
pedicellate. Ascospores (9.5–)11–13(–16) × 5.5–7 µm
(x̄ = 12.5 × 6.5µm, n = 50), uni-seriate, ellipsoidal to broad-
fusiform or triangular, aseptate, hyaline to pale brown,
smooth-walled, with guttules and an apical germ pore.
Asexual morph (associated): Hyphomycetous. Conidi-
ophores (32.5–)42–55(–95) × 3.5–14 µm (x̄ = 53 × 7µm,
n = 10), arising from the substrate, macronematous, mon-
onematous, cylindrical, sparingly branched, pale brown
to brown, septate, smooth-walled. Conidiogenous cells
(10.5–)14–20(–22) × 3–7.5µm (x̄ = 17.5 × 5.5µm, n = 10),
terminal or intercalary, polytretic, brown, tapering, apex
rounded, conidia formed through minute pores. Conidia
(10–)17–20(–25) × 6–8µm (x̄ = 18 × 7µm, n = 30), solitary,
ellipsoidal to subglobose, 0–1-septate, sometimes con-
stricted at septum, slightly curved, smooth-walled, hyaline
to brown, with guttules.
Fungal Diversity
1 3
Material examined: Sweden, Uppland, Fresta, on Cla-
vulina cinerea, 18 September 1984, von H. Eichwald (S-F
135699); Sweden, Södermanland, Hölö sn. Tullgarnsomra-
det, SSO, Tullgarns slot, on Clavulina sp., 9 October 2008,
K. Jaederfeldt (S-F 121831).
Known hosts and distribution: On Clavaria fuliginea in
Franch (type locality) (Desmazières 1837); on Clavaria in
Franch, Germany, Sweden, UK and USA (Samuels etal. 1997).
Notes: Helminthosphaeria clavariarum is fungicolous and
has been reported on Clavariaceae members in Europe and
North America associated with diplococcium-like asexual
morphs (Dennis 1978; Samuels etal. 1997). Huhndorf etal.
(2004b) analyzed LSU sequence data for H. clavariarum
(SMH4609). Miller etal. (2014) re-examined the material
of SMH4609 and analyzed TUB sequence data for H. cla-
variarum (ANM Acc.17). In this study, we re-examined two
authentic specimens (F 135699 and F 121831) determined
by Lundqvist (mentioned in the label of material).
New combination:
Helminthosphaeria plumbea (Huhndorf, F.A. Fernández &
Cand.) S.K. Huang & K.D. Hyde, comb. nov.
Fig. 23 Helminthosphaeria
clavariarum: a–s (S-F 135699);
t–cc (S-F 121831). a Host of
Clavulina cinerea. b Ascomata
on host. c, e Ascomata in cross
section. d Setae. f Peridium
with cylindrical cells of ostiole
in the upper part. g Asci with
paraphyses (arrows indicate a
cell on ascogenous hyphae). h
Septate paraphyses. i, j Asci.
k–s Ascospores. t Colonies on
host. u, v Conidiophores and
conidia. w–z, aa–cc Conidia.
Scale bars: b = 500µm,
c = 200µm, d–e = 100µm,
f = 50µm, g–j. u. v = 20µm,
k–s. w–z. aa–cc = 10µm
Fungal Diversity
1 3
Basionym: Synaptospora plumbea Huhndorf, F.A.
Fernández & Cand., Sydowia 51(2): 177 (1999)
Index Fungorum number: IF558251
Notes: See notes for Synaptospora.
Hilberina Huhndorf & A.N. Mill., Mycol. Res. 108(1): 31
(2004)
Saprobic on wood or leaves. Sexual morph: Ascomata
perithecial, solitary or scattered, superficial, membrana-
ceous, ovoid to obpyriform, brown to dark brown, tubercu-
late, covered with setae, ostiolate. Asci 8-spored, unitunicate,
cylindrical. Ascospores biseriate, cylindrical, geniculate, one
or both end(s) tapering to a distinct point or rounded ends,
hyaline when young, becoming pale brown, smooth-walled,
with guttules, 0–3-septate, sometimes with sheath. Asexual
morph: Hyphomycetous. Conidiophores macronematous,
mononematous, cylindrical, single or in group up to two,
erect, arising directly from the host, multi-septate, cells
expanding sometimes, brown to dark brown, apical cell
swollen. Conidiogenous cells phialides, with cylindrical
branches, hyaline to pale brown, discrete, smooth-walled;
secondary branches, hyaline, 2–3 per branch, cylindrical,
tapering at the apex. Conidia hyaline, aseptate, oblong to
obovoid, smooth-walled (adapted from Fuckel 1870; Miller
and Huhndorf 2004a).
Notes: Fuckel (1870) introduced Leptospora cau-
data has black ascomata and oblong ascospores and was
accompanied with a hyphomycete with hyaline, oval
conidia, on decayed wood of Betula alba in nature. This
species was introduced as the generic type of Hilberina
(Miller and Huhndorf 2004a). Subsequently, several taxa
with bristly ascomata and ascospores distinctly curved or
bent near the attenuate, lanceolate or acerose base were
accepted as members of Hiberina (Declercq 2009, 2013;
Miller etal. 2014). Miller etal. (2014) sequenced five
Hilberina species and found that they are polyphyletic
in Helminthosphaeriaceae and their asexual morphs are
unknown. In this study, Hilberina species are closely
related to members of Helminthosphaeria and scattered in
Helminthosphaeriaceae (Fig.21). Its sexual-asexual link
needs confirming with cultural characteristics and more
phylogenetic data.
Type species: Hilberina caudata (Fuckel) Huhndorf & A.N.
Mill., in Miller & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Basionym: Leptospora caudata Fuckel, Jb. nassau. Ver.
Naturk. 23-24: 144 (1870)
Facesoffungi number: FoF 10013; Fig.24
Saprobic on decorticated wood. Sexual morph: Asco-
mata 350–600 × 375–565 µm (x̄ = 475 µm × 475 µm,
n = 10), perithecial, solitary or scattered, superficial, mem-
branaceous, ovoid to subglobose, dark brown to black, cov-
ered with setae, ostiolate, with papilla, periphysate. Setae
15–30µm wide at base, brown to pale brown, tapering,
with a septate lumen and apex rounded. Peridium 35–80µm
(x̄ = 55µm, n = 30) wide, comprising two layers, outer
layer composed of pale brown to brown cells of textura
angularis; inner layer composed of hyaline cells of textura
prismatica. Asci 125–180 × 9.5–13.5µm, 8-spored, unitu-
nicate, cylindrical, pedicellate, apex rounded, with J- incon-
spicuous apical ring. Ascospores (38–)43–50 × 3–8 µm
(x̄ = 45.5 × 5.5µm, n = 50), biseriate, cylindrical, genicu-
late, one end tapering to a distinct point, hyaline when
young, becoming pale brown, smooth-walled, with gut-
tules, 0–3-septate, without gelatinous appendages. Asex-
ual morph (associated): Hyphomycetous. Conidiophores
3–8.5µm, cylindrical, single or in group up to two, erect,
arising directly from the host, macronematous, mononema-
tous, multi-septate, not constricted, cells expanding some-
times, brown to dark brown, apical cell swollen. Conid-
iogenous cells phialides (3–)5–8 × 1–2µm (x̄ = 6 × 1.5µm,
n = 30), with cylindrical branches, hyaline to pale brown,
discrete, smooth-walled, aseptate; secondary branches
(3.5–)4.5–5.5(–7.5) × 1–1.5µm (x̄ = 5 × 1.2 µm, n = 30),
hyaline, aseptate, 2–3 per branch, cylindrical, tapering at
the apex. Conidia 2.5–3.5 × 1–2µm (x̄ = 3 × 1.5µm, n = 50),
hyaline, aseptate, oblong to obovoid, smooth-walled
(adapted from Fuckel 1870).
Material examined: Puerto Rico, Bisley Watershed 3,
down slope on Quebrada 2 to Quebrada 3, Luquillo Moun-
tains, 18º 19.00′ N/65° 0.07′ W, on dead wood, 28 January
1997, F.A. Fernández (F-SMH 3156); Germany, Hessen, 9º
0′ 19″ E/49° 46′ 28″ N, on dead wood of Betula sp., K.W.G.
Fuckel (G-G00266357 syntype).
Known hosts and distribution: On decorticated wood of
Betula alba in Germany (type locality) (Fuckel 1870); on
dead wood in Puerto Rico.
Notes: In this study, we re-examined a typical material
and an authentic specimen, which were well preserved.
The sexual morph of Hilberina caudata associated with
haplographium-like asexual morph were found on the type
materials. Hilberina caudata is sister to Hilberina punctata
and nested in Helminthosphaeriaceae based on multi-gene
analysis (Miller etal. 2014; this study, 96%ML/1.00PP,
Fig.21). However, cultural characteristics and phylogenetic
data of H. caudata are required to gain better insights into
the relationship with Haplographium and its taxonomic
placement.
Ruzenia O. Hilber, The Genus Lasiosphaeria and Allied
Taxa (Kelheim): 7 (2002)
Saprobic on wood. Sexual morph: Ascomata perithecial,
scattered to gregarious, superficial, subglobose to obovoid,
dark brown to black, rough, ostiolate, with papilla, peri-
physate. Peridium membranaceous, composed of brown to
hyaline cells of textura angularis to textura prismatica. Asci
Fungal Diversity
1 3
8-spored, unitunicate, cylindrical, pedicellate, apex rounded,
with J- apical ring. Ascospores allantoid, 0–1-septate, hya-
line, smooth-walled, guttulate. Asexual morph: Undeter-
mined (adapted from Hilber and Hilber 2002).
Notes: The monotypic genus Ruzenia was introduced as
similar to Lasiosphaeria (Hilber and Hilber 2002). Subse-
quently, its generic type, Ruzenia spermoides, was clarified
as member of Helminthosphaeriaceae based on phylogenetic
analyses (Miller and Huhndorf 2004a; Miller etal. 2014).
In this study, Ruzenia is similar to Hilberina, but differs
in having less ascomata setae, lacking basal spines on the
ascospores and being hyaline. It is basal to Helminthospha-
eria plumbea and Hilberina munkii in Helminthospha-
eriaceae (90%ML/0.93PP, Fig.21).
Type species: Ruzenia spermoides (Hoffm.: Fr.) O. Hil-
ber ex A. N. Mill. & Huhndorf, Mycological Research 108
(1): 31 (2004)
Basionym: Sphaeria spermoides Hoffm., Veg. Crypt. 2:
12 (1790)
Facesoffungi number: FoF 10014; Fig.25
Fig. 24 Hilberina caudata:
a, c, h, m–p (G-G00266357,
Syntype); b, d–g, j–l (F-SMH
3156); i (redrawn from Fuckel
1870). a Scattered ascomata. b
Ascoma on host. c Ascoma in
longitudinal section surrounded
by setae. d Ascoma in cross
section. e Peridium. f Setae.
g–i Asci. j–l Ascospores. m
Conidiophores with conidia.
n, o Primary and secondary
branches of conidiogenous cells
with conidia. p Conidia. Scale
bars: a = 1mm, b. c = 500µm,
d = 200µm, e. h. i. m = 50µm,
f = 100µm, g. j–l. n. o. = 20µm,
p = 10µm
Fungal Diversity
1 3
Saprobic on wood. Sexual morph: Ascomata
385–620 × 360–530µm (x̄ = 485 × 465µm, n = 10), per i-
thecial, scattered to gregarious, superficial, subglobose to
obovoid, dark brown to black, rough, ostiolate, with papilla,
periphysate. Peridium 75–105µm (x̄ = 90 µm, n = 30)
wide, membranaceous, comprising two layers, outer layer
composed of brown cells of textura angularis; inner layer
composed of hyaline cells of textura prismatica. Para-
physes 3–7.5µm wide, septate, constricted at the septum,
unbranched, cylindrical, tapering towards the rounded apex.
Asci (100–)110–130(–150) × 8–12 µm (x̄ = 125 × 10 µm,
n = 30), 8-spored, unitunicate, cylindrical, pedicellate, apex
rounded, with J- apical ring. Ascospores 20–26 × 3–5.5µm
(x̄ = 23 × 4 µm, n = 50), bi-seriate, allantoid, 0–1-sep-
tate, hyaline, smooth-walled, guttulate. Asexual morph:
Undetermined.
Material examined: Sweden, Uppland, Bladaker; on dead,
decorticated wood of Tilia cordata, 20 September 1992, N.
Lundqvist (S-F 242341); Sweden, Uppland, Österaker-Östra
Ryd, on dead, decorticated wood, 17 May 1983, N. Lun-
dqvist (S-F 242342).
Known hosts and distribution: On dead decorticated
wood in Czech Republic, Denmark, Germany (type local-
ity), Sweden and USA (Hoffmann 1790; Fries 1823a; Miller
and Huhndorf 2004a).
Notes: Ruzenia spermoides strains clustered in Hel-
minthosphaeriaceae with 100%ML/1.00PP support and this
clade is basal to Helminthosphaeria plumbea and Hilberina
munkii (90%ML/0.93PP, Fig.21) in the phylogenetic analy-
sis (Miller and Huhndorf 2004a; Miller etal. 2014). In this
study, we were unable to obtain the type material. Therefore,
we re-examined two authentic samples collected by Lun-
dqvist in Sweden.
Sordariales Chadef. ex D. Hawksw. & O.E. Erikss., Syst.
Ascom. 5(1): 182 (1986)
Sordariales comprises Bombardiaceae, Chaetomiaceae,
Diplogelasinosporaceae, Lasiosphaeriaceae, Lasiosphaeri-
daceae, Naviculisporaceae, Podosporaceae, Neoschizothe-
ciaceae, Strattoniaceae, Sordariaceae and Zygospermel-
laceae as clarified by phylogenetic analyses (Hawksworth
and Eriksson 1986; Huhndorf etal. 2004b; Kruys etal.
Fig. 25 Ruzenia spermoides:
a–c, e, f, j, k (S-F242341);
d, g–i, m, n (S-F242342).
a Gregarious ascomata. b
Ascomata on host. c, d Ascoma
in cross section. e Peridium.
f Asci with paraphyses. g–i.
Asci. j–n Ascospores. Scale
bars: b = 500µm, c. d = 200µm,
e. g–i = 50µm, f = 100µm,
j–n = 10µm
Fungal Diversity
1 3
2014; Maharachchikumbura etal. 2016; Hongsanan etal.
2017; Hyde etal. 2017, 2020; Marin-Felix etal. 2020; this
study). Most taxa of Sordariales occur in terrestrial or/and
aquatic habitats as saprobes and have ascospores with vari-
ous appendages or different wall patterns (Maharachchi-
kumbura etal. 2015, 2016; Hyde etal. 2020). This order is
basal to Phyllachorales and Boliniales in Sordariomycetidae
(59%ML, Fig.1).
Bombardiaceae S.K. Huang, Maharachch. & K.D. Hyde,
fam. nov.
Facesoffungi number: FoF 10141; Index Fungorum num-
ber: IF558344
Etymology: Named after Bombardia, the oldest genus in
this family.
Coprophilous or saprobic on wood or isolated from soil.
Sexual morph: Ascomata perithecial or cleistothecial, soli-
tary to scattered or gregarious, immersed to semi-immersed
to superficial, globose to subglobose, glabrous or hairy,
yellow to dark brown to black, sometimes semitransparent,
membranaceous or coriaceous. Paraphyses numerous, fili-
form to cylindrical, septate, evanescent. Asci 4–8-spored,
unitunicate, cylindrical to clavate, pedicellate, usually with
J- apical ring, sometimes with apical globule. Ascospores
uni-seriate or bi-seriate, ellipsoidal or cylindrical or maraca-
shaped, aseptate to multi-septate, hyaline to brown to dark
brown, verrucose or smooth-walled, sometimes with a gelat-
inous sheath or irregular appendage(s). Asexual morph:
Hyphomycetous. Conidiophores arising from aerial hyphae,
erect, inflated at the base. Phialides monophialidic, smooth-
walled, hyaline, cylindrical, unbranched or branched.
Conidia small, hyaline to pale yellow, ellipsoidal to oval,
aseptate, smooth-walled, forming globose masses on the
apex of the phialides (adapted from Malloch and Cain 1971;
Lundqvist 1964a, b, 1972; Udagawa and Furuya 1974; Khan
and Krug 1991; Krug and Scott 1994; Bell 2005; Davison
etal. 2008).
Type genus: Bombardia (Fr.) P. Karst. 1873
Notes: Marin-Felix etal. (2020) mentioned that Apodos-
pora, Bombardia, Bombardioidea, Fimetariella and several
species of Cercophora, Podospora and Zopfiella formed
a low support clade. In this study, these taxa clustered
(75%ML) and is sister to Sordariaceae (74%ML, Fig.26).
The species of this cluster have diverse morphological
characteristics. Most of them are coprophilous fungi, except
for Zopfiella attenuata and Ramophialophora vesiculosa col-
lected from soil and, Bombardia bombarda found on hard-
wood (Lundqvist 1972; Udagawa and Furuya 1974). They
have hairy or glabrous, membranaceous (sometimes with
carbonaceous neck), or coriaceous ascomata (Malloch and
Cain 1971; Lundqvist 1972; Udagawa and Furuya 1974;
Khan and Krug 1991; Krug and Scott 1994; Bell 2005;
Davison etal. 2008). The hyphomycetous R. vesiculosa has
globose to ellipsoidal conidia with a truncate base (Calduch
etal. 2004). Zopfiella attenuata and Z. pleuropora have cle-
istothecial ascomata and ascospores comprise fusiform to
ellipsoidal, brown upper cell(s) and a conical hyaline lower
cell, without appendages or a sheath (Malloch and Cain
1971; Udagawa and Furuya 1974); the other members have
perithecial ascomata and ascospores have appendages or
surround the gelatinous sheath (Lundqvist 1964a, b, 1972;
Khan and Krug 1991; Krug and Scott 1994; Krug 1995;
Bell 2005; Davison etal. 2008). Apodospora peruviana, A.
simulans, A. gotlandica, Bombardioidea anartia and Fim-
eteriella rabenhorstii have broadly fusiform to ellipsoidal
ascospores surrounded by a gelatinous sheath (Lundqvist
1964a, b, 1972; Krug and Scott 1994; Krug 1995; Davison
etal. 2008). All Apodospora species have the gelatinous
sheath with an apical invagination (Lundqvist 1972; Krug
and Scott 1994; Krug 1995; Davison etal. 2008). Fime-
teriella rabenhorstii is the only species in the cluster with
4-spored asci (Lundqvist 1964a, b; Krug 1995). Bombar-
dioidea anartia and Podospora appendiculata have a spe-
cial coriaceous peridium similar to Bombardia bombarda
(Lundqvist 1972; Krug and Scott 1994). Whereas, Bombar-
dia bombardia, Cercophora scortea, Podospora fabiformis,
P. fibrinocaudata and P. appendiculata have ascospores
comprise an ellipsoidal, brown upper cell and a cylindri-
cal, hyaline pedicel, with terminal appendage(s) at each or
one end (Lundqvist 1972; Khan and Krug 1991; Bell 2005).
Podospora fabiformis has ascospores with apical and basal
appendages, sometimes two lateral appendages arising from
the top of the pedicel (Bell 2005). Podospora fibrinocau-
data has ascospores with a gelatinous sheath surrounding
the pedicel’s basal (Khan and Krug 1991). Bombardia bom-
barda, Cercophora scortea and Podospora appendiculata
have cylindrical to clavate asci with a subapical or apical
globule and ascospores with appendages at each end (Lun-
dqvist 1972).
Apodospora Cain & J.H. Mirza, Can. J. Bot. 48(5): 891
(1970)
Coprophilous or saprobic on wood. Sexual morph:
Ascomata perithecial, gregarious to scattered, immersed to
semi-immersed, globose to subglobose, dark brown to black,
glabrous or hairy. Peridium membranaceous, composed of
hyaline to brown cells of textura angularis to textura pris-
matica. Paraphyses numerous, filiform, septate, evanes-
cent. Asci 8-spored, unitunicate, ellipsoidal, pedicellate,
blunt at the apex. Ascospores oval to ellipsoidal, aseptate,
brown to dark brown, verrucose, surrounded by gelatinous
sheath, usually with an apical invagination. Asexual morph:
Hyphomycetous. Mycelium grey-brown, with numerous
phialides scattered on hyphae. Conidia small, ovate masses
on the apex of the phialides (adapted from Cain and Mirza
1969).
Fungal Diversity
1 3
Fig. 26 Phylogram generated
from maximum likelihood anal-
ysis based on combined LSU,
ITS, TUB and RPB2 sequence
data of Sordariales. The confi-
dence 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 correspond-
ing nodes. The ‘--’ indicates
lack of statistical support
(< 50% for ML-BS and < 0.90
for BY-PP). Two hundred and
thirty-five strains are included
in the combined analyses,
which comprise 2877 characters
(851 characters for LSU, 509
characters for ITS, 665 char-
acters for TUB, 852 characters
for RPB2) after alignment.
Strains of Microascales are
used as outgroup taxa. The best
score in IQ-TREE explores
with a final likelihood value
of −83459.1151 is presented.
The model of each partitioned
gene is: LSU: GTR + F + I + G4;
ITS: TIM2e + I + G4; TUB:
HKY + F + I + G4; RPB2:
TIM3 + F + I + G4. Strain
numbers are noted after the spe-
cies names and ex-type strains
are marked with ‘T’ after the
culture number. Alignments are
available at TreeBASE (URL:
http:// purl. org/ phylo/ treeb ase/
phylo ws/ study/ TB2: S28270)
Strattoniaceae
Chaetomiaceae
Podosporaceae
Diplogelasinosporaceae
Sordariales
TriangulariaCladorrhinumPodospora
T. praecox
T.tarvisina
T.nannopodalis
T.arizonensis
T.microsclerotigena
T.yaeyamensis
T.striata
T.samala
T.tetraspora
T.unicaudata
C.leucotrichum
C.terricola
C.tomentosum
C.grandiusculum
C.olerum
C.coprophilum
P.sacchari
P.brunnescens
P.striatispora
P.macrospora
P.costaricensis
P.dacryoidea
P.flexuosa
P.hamata
P.jamaicensis
P.spinosa
S.petrogale
Strattonia
Podospora fimicola CBS 990.96
Chrysanthotrichum peruvianum CBS 732.68 T
Brachychaeta variospora CBS 414.73 T
Podospora petrogale CBS:109409 T
Cercophora samala CBS:307.81 T
Triangularia batistae CBS 381.68 T
Podospora praecox CBS 251.71 T
Triangularia pauciseta CBS 451.62
Madurella tropicana CBS 201.38 T
Thermothelomyces guttulata CGMCC:3.15185 T
Crassicarpon hotsonii CBS:389.93 T
Chaetomium globosumCBS 160.62 T
Canariomyces subthermophilus CBS 509.74 T
Chrysocorona lucknowensis CBS 727.71 T
Dichotomopilus funicola CBS 159.52 T
Collariella robusta CBS:551.83 T
Triangularia phialophoroides CBS 301.90 T
Corynascus sepedonium CBS 111.69 T
Floropilus chiversii CBS 558.80 T
Arnium olerum CBS 120012
Cladorrhinum intermedium CBS 433.96 T
Arcopilus cupreus CBS 560.80
Apiosordaria striatispora CBS 154.77 T
Canariomyces notabilis CBS 548.83 T
Triangularia bambusae CBS 352.33 T
Cladorrhinum brunnescens CBS:643.75A T
Cercophora striata SMH3431
Thermothelomyces thermophila CBS 381.97
Thermothelomyces heterothallica CBS 202.75 T
Corynascus novoguineensis CBS:359.72 T
Podospora fimicola CBS 482.64 T
Triangularia backusii CBS 539.89 T
Podospora nannopodalis CBS 113680
Podospora unicaudata CBS 313.58 T
Parathielavia appendiculata CBS 723.68 T
Subramaniula thielavioides CBS:122.78 T
Thermothielavioides terrestris CBS 117535 T
Collariella carteri CBS 128.85 T
Ovatospora pseudomollicella CBS:251.75 T
Diplogelasinospora moalensis CBS 136018
Subramaniula obscura CBS:132916 T
Achaetomium luteum CBS 544.83
Acrophialophora fusispora CBS 380.55 T
Triangularia ovina CBS 671.82 T
Condenascus tortuosus CBS 610.97
Diplogelasinospora inaequalis CBS:436.74 T
Achaetomium lippiae URM:7547 T
Podospora tarvisina CBS 265.70
Arcopilus fusiformis CBS 484.85
Trichocladium gilmaniellae CBS 388.75 T
Strattonia oblecythiformis CBS:110350 T
Canariomyces vonarxii CBS 160.80 T
Ovatospora medusarum CBS:148.67 T
Apiosordaria spinosa CBS 259.71 T
Microthielavia ovispora CBS 165.75 T
Triangularia allahabadensis CBS 724.68 T
Trichocladium asperum CBS 903.85 T
Diplogelasinospora princeps FMR 13415
Apiosordaria jamaicensis CBS 672.70 T
Amesia gelasinospora CBS 673.80 T
Podospora inflatula CBS 413.82
Parathielavia hyrcaniae CBS 353.62 T
Carteria arctostaphyli CBS 229.82 T
Arnium tomentosum Francoise Candoussau
Chaetomium tenue CBS 139.38 T
Cercophora costaricensis SMH4021
Podospora bulbillosa CBS 304.90 T
Thermothielavioides terrestris CBS 492.74
Zopfiella leucotricha CBS 463.61
Pseudothielavia subhyaloderma CBS 473.86 T
Pseudothielavia arxii CBS 603.97 T
Dichotomopilus variostiolatus CBS:179.84 T
Arnium arizonense CBS 120289
Amesia atrobrunnea CBS 379.66 T
Trichocladium griseum CBS 119.14
Cercophora terricola ATCC 200395
Cladorrhinum australe INTA-AR:70 T
Corynascus citrinus BCC:79098 T
Acrophialophora nainiana CBS 100.60 T
Stolonocarpus gigasporus CBS 112062 T
Apiosordaria sacchariCBS 713.70 T
Parathielavia kuwaitensis CBS 945.72 T
Madurella mycetomatis CBS 109801 T
Apiosordaria hamata CGMCC 3.15230 T
Zopfiella macrospora CBS 286.86 T
Cladorrhinum microsclerotigenum CBS:290.75 T
Chaetomium pseudoglobosum CBS 574.71 T
Madurella pseudomycetomatis CBS 129177 T
Triangularia setosa CBS 311.58
Crassicarpon thermophilum CBS:406.69 T
Apiosordaria yaeyamensis NBRC 31170 T
Cladorrhinum flexuosum CBS:126090 T
Amesia nigricolor CBS 600.66 T
Subramaniula asteroides CBS:123294 T
Acrophialophora hechuanensisGZUIFR-H08-1 T
Zopfiella tetraspora CBS 245.71 T
Stellatospora terricola CBS 811.95 T
Triangularia anserina CBS 433.50
Cladorrhinum globisporum LC5415 T
Cladorrhinum foecundissimum CBS 180.66 T
Triangularia longicaudata CBS 252.57 T
Collariella causiiformis CBS 792.83 T
Chrysanthotrichum lentum CBS 339.67 T
Pseudothielavia terricola CBS 165.88 T
Chrysanthotrichum leptolentum CBS 126.85 T
Ovatospora unipora CBS 109.83 T
Arcopilus flavigenus CBS 337.67 T
Cercophora grandiuscula CBS 120013
Hyalosphaerella fragilis CBS 456.73 T
Cercophora coprophilaSMH3794
Triangularia verruculosa CBS 148.77
Cladorrhinum hyalocarpum CBS 322.70 T
Dichotomopilussubfunicola CGMCC 3.12892 T
Diplogelasinospora grovesii CBS:340.73 T
Achaetomium macrosporum CBS 152.97 T
100/1.00
71/0.96
64/--
100/1.00
100/1.00
100/1.00
95/--
66/--
100/1.00
100/0.96
100/0.97
63/--
100/1.00
51/--
100/1.00
98/1.00
69/--
94/0.97
100/1.00
100/1.00
61/--
100/1.00
52/--
100/1.00
100/1.00
100/1.00
100/1.00
98/--
100/1.00
88/0.99
100/1.00
62/--
99/--
90/1.00
89/0.92
98/0.99
99/--
100/1.00
97/0.96
100/1.00
98/0.92
100/1.00
100/1.00
68/--
100/1.00
66/--
100/1.00
100/1.00
95/0.95
100/1.00
100/1.00
100/1.00
100/1.00
100/1.00
83/0.98
100/1.00
100/0.98
94/--
100/1.00
73/--
93/--
100/1.00
98/1.00
100/1.00
83/1.00
100/1.00
65/1.00
89/--
100/1.00
100/1.00
100/1.00
99/1.00
74/--
100/0.99
95/0.93
100/1.00
100/1.00
93/0.99
100/1.00
100/1.00
80/--
53/--
100/1.00
98/1.00
100/1.00
100/1.00
87/1.00
98/--
100/1.00
99/1.00
90/0.90
100/1.00
80/--
100/1.00
98/0.95
100/0.98
100/1.00
97/0.99
100/1.00
71/--
99/1.00
85/--
58/--
100/1.00
100/1.00
68/--
100/1.00 81/--
99/1.00
100/1.00
89/--
Fungal Diversity
1 3
Podospora didyma CBS 232.78
Podospora communis CBS 118393
Schizothecium aloides CBS 879.72
Boothiella tetraspora CBS 334.67 T
Cercophora scortea GJS L556
Jugulospora rotula FMR 12781
Pseudoneurospora canariensis CBS 135818 T
Zopfiella attenuata CBS:266.77 T
Schizothecium curvisporum CBS:507.50 T
Neurospora tetrasperma CBS 223.38
Lasiosphaeria lanuginosa SMH3819
Apodus deciduus CBS 506.70 T
Apodospora peruviana CBS 118394
Berkeleyomyces rouxiae CBS 178.82
Ramophialophora humicola CBS:124563 T
Anopodium ampullaceum MJR 40/07
Cercophora sulphurella SMH2531
Fimetariella rabenhorstii Lundqvist 20410-c
Bombardia bombarda SMH4821
Cercophora appalachianensis HKUCC3711
Apodus oryzae CBS 376.74
Rhypophila decipiens CBS 258.69
Cercophora solaris ANM 734
Schizothecium conicum CBS 434.50
Lasiosphaeria ovina SMH 1538
Cercophora thailandica MFLUCC:12-0845 T
Cercophora aquatica JF 09214 T
Lasiosphaeris hirsuta JF02183
Neurospora crassa ICMP 6360
Immersiella immersa SMH4104
Zopfiella tardifaciens CBS 670.82 T
Areotheca areolata UAMH7495
Arnium japonense SANK 10273
Sordaria nodulifera NBRC 32551 T
Thielavia basicola CBS 178.82 T
Zopfiella submersa CBS 698.96 T
Zopfiella pilifera CBS 413.73 T
Episternus onthophagi KRAM:F58223 T
Areotheca ambigua CBS 215.60
Zygospermella insignis E00204312
Zopfiella marina CBS 155.77 T
Rinaldiella pentagonospora CBS 132344 T
Zopfiella erostrata CBS 255.71
Podospora prethopodalis CBS 121128
Apiosordaria microcarpa CBS 692.82 T
Schizothecium selenosporum CBS 109403 T
Lasiosphaeris hispida SMH3336
Lasiosphaeria similisorbina AR:1884 T
Echria gigantospora ATCC:MYA:4170 T
Lundqvistomyces tanzaniensis TRTC51981
Microthecium tenuissimum CBS 112764 T
Echria macrotheca Lundqvist 2311
Ramophialophora globispora LC6218
Zopfiella pleuropora CBS:518.70 T
Pseudoechria prolifica CBS 250.71 T
Podospora fabiformis CBS 112043 T
Sordaria tamaensis NBRC 32552 T
Bombardioidea anartia HHB99-1
Zygopleurage zygospora SMH4219
Jugulospora carbonaria ATCC 34567
Lasiosphaeria rugulosa ATCC:MYA:2578 T
Microthecium quadrangulatum CBS 112763 T
Immersiella immersa SMH2589
Microthecium retisporum CBS 995.72
Apodospora gotlandica E00204952
Microthecium zobelii CBS 268.62
Cercophora sparsa JF00229
Cercophora arenicola ANM 1080
Podospora serotina CBS 252.71
Corylomyces selenosporus CBS 113930 T
Pseudoechria longicollis CBS 368.52 T
Schizothecium inaequale CBS 356.49 T
Schizothecium fimbriatum CBS 144.54
Boothiella tetraspora CBS 887.97
Lasiosphaeria glabrata SMH4617
Cercophora newfieldiana SMH3303
Cercophora mirabilis CBS 120402
Podospora minicauda CBS 227.87
Arnium mendax E00122117
Lasiosphaeris hirsuta SMH1543
Ramophialophora vesiculosa CBS 110629 T
Apodospora simulans Kruys 701
Berkeleyomyces basicola CBS 341.33
Pseudoechria curvicolla CBS 368.69
Schizothecium carpinicola CBS:228.87 T
Neurospora metzenbergii FGSC 8847
Pseudoechria decidua CBS 254.71 T
Mammaria echinobotryoides CBS 458.65
Lasiosphaeris hispida CBS 955.72
Arnium cirriferum CBS 120041
Podospora appendiculata CBS 212.97
Neurospora hispaniola FGSC 8817 T
Sordaria islandica CBS 512.77 T
Lasiosphaeria sorbina CBS 885.85
Podospora excentrica CBS 118392
Pseudoschizothecium atropurpureum SMH3073
Schizothecium tetrasporum CBS 394.87
Neurospora sitophila CBS 112.19
Lundqvistomyces karachiensis CBS 657.74
Mammaria echinobotryoides CBS 277.63
Lasiosphaeria ovina CBS 126299
Podospora bullata CBS:115576 T
Immersiella caudata SMH3298
Lasiosphaeria miniovina SMH:2392 T
Strattonia minor CBS 380.86
Apiosordaria vermicularis CBS 303.81 T
Schizothecium glutinans CBS 134.83
Triangularia mangenotii CBS:419.67 T
Microthecium fimicola CBS 967.97
Pseudoneurospora amorphoporcata CBS 626.80
Ramophialophora petraea LC6222
Rhypophila cochleariformis CBS 249.71
Podospora cupiformis CBS:246.71 T
Jugulospora rotula FMR 12690
Podospora intestinacea CBS 113106
Podospora leporina CBS 365.69
Podospora fibrinocaudata CBS 315.91 T
Zopfiella latipes IFO 9826
Zopfiella tabulata CBS 230.78
Bellojisia rhynchostoma CBS 118484
Arnium hirtum E00204950
0.2
74/--
59/0.98
94/--
100/0.96
100/1.00
84/--
50/0.96
100/1.00
72/--
84/1.00
88/--
100/1.00
99/1.00
100/1.00
100/0.98
88/--
100/1.00
100/1.00
98/--
96/--
100/0.97
69/--
83/0.99
99/1.00
100/--
100/--
100
59/--
100/1.00
99/--
75/0.99
61/--
99/0.99
95/--
57/0.97
100/1.00
100/1.00
75/--
100/1.00
98/--
92/--
100/1.00
100/1.00
99/0.95
100/1.00
52/--
100/1.00
86/--
100/1.00
50/--
100/1.00
100/1.00
96/--
100/1.00
54/0.97
100/1.00
100/1.00
100/1.00
95/--
100/1.00
100/1.00
94/--
100/1.00
100/1.00
83/1.00
100/1.00
100/1.00
92/0.96
100/1.00
96/--
100/1.00
100/1.00
100/1.00
99/0.97
100/1.00
66/--
100/--
100/1.00
99/1.00
100/1.00
90/--
56/--
89/--
100/0.95
99/--
99/--
100/1.00
100/1.00
99/1.00
90/0.99
100/1.00
56/1.00
74/--
74/--
92/0.92
100/--
67/--
100/1.00
99/1.00
100/--
100/1.00
100/0.96
100/0.93
100/1.00
62/--
99/1.00
100/1.00
92/--
100/1.00
64/--
100/1.00
83/--
100/1.00
88/--
90/--
100/1.00
98/--
100/1.00
100/0.99
100/0.96
Jugulospora
Immersiella
Neoschizothecium
Lasiosphaeris
Microascales
Lasiosphaeridaceae
Neoschizotheciaceae
Zygospermellaceae
Mammaria echinobotryoides
Sordariaceae
Naviculisporaceae
Sordariales
Lasiosphaeriaceae
Bombardiaceae
Clade I
J. minor
I. hirta
N. inaequale
N. tetrasporum
N. curvisporum
N. selenosporum
N. fimbriatum
N. glutinans
N. minicaudum
N. aloides
N. carpinicola
N. conicum
L. arenicola
Fig. 26 (continued)
Fungal Diversity
1 3
Notes: Cain and Mirza (1969) introduced Apodospora typi-
fied by A. simulans, which has oblong to ellipsoidal ascospores
surrounded by a gelatinous sheath and oval to globose small
conidia masses on the phialides. This genus resembles Sord-
aria in having ascospores surrounded by a gelatinous sheath
(Cain and Mirza 1969). Six species are accommodated in this
genus (Cain and Mirza 1969; Lundqvist 1972; Fakirova 1973;
Muroi etal. 1987), and A. gotlandica, A. peruviana, and A.
simulans have molecular data (Kruys etal. 2014). In this study,
Apodospora species clustered in Bombardiaceae in the phylo-
genetic analysis (100%ML/1.00PP, Fig.26).
Type species: Apodospora simulans Cain & J.H. Mirza,
Can. J. Bot. 48(5): 891 (1970)
Facesoffungi number: FoF 10015; Fig.27
Coprophilous. Sexual morph: Ascomata
380–530 × 250–400µm (x̄ = 410 × 275µm, n = 5), perithe-
cial, gregarious to scattered, immersed to semi-immersed,
globose to subglobose, dark brown to black, ostiolate,
with papilla. Peridium 25–50µm (x̄ = 38µm, n = 30) wide,
membranaceous, comprising two layers, outer layer com-
posed of pale brown to brown cells of textura angularis;
inner layer composed of hyaline cells of textura prismatica.
Paraphyses 3–4µm wide, filiform, septate, evanescent.
Asci (200–)220–280(–350) × 12–20µm (x̄ = 250 × 16µm,
n = 30), 8-spored, unitunicate, ellipsoid, pedicellate,
apex blunt, with J- apical ring distinct. Ascospores
(15–)18–21(–25) × 8–10.5µm (x̄ = 20 × 9µm, n = 50), uni-
seriate, oblong to oval, aseptate, brown to dark brown, ver-
rucose, surrounded by a gelatinous sheath 2.5–6µm diam.
Asexual morph: Hyphomycetous. Mycelium grey-brown,
with numerous phialides scattered on the hyphae. Phi-
alides pale brown, with flaring collarette. Conidia small,
ovate forming globose masses on the apex of the phialides
(adapted from Cain and Mirza 1969).
Fig. 27 Apodospora simu-
lans: a (redrawn from Cain
and Mirza 1969); b–k (S-F
122,911). a Conidophores with
conidia. b Herbarium material.
c Ascomata on host. d Ascoma
in cross section. e Peridium. f,
g Asci (enlarged: apical ring).
h–k Ascospores. Notes: Fig
d–k are stained in KOH. Scale
bars: a = 10µm, c = 500µm,
d = 200µm, e–g = 50µm,
h–k = 20µm
Fungal Diversity
1 3
Material examined: Sweden, Uppland, Danmark par.,
500m E. of Grynbacken, in coniferous forest, on old moose
dung, 18 May 1959, N. Lundqvist (S-F122911).
Known hosts and distribution: On dung of moose (Alces
americanus) in Canada (type locality) (Cain and Mirza
1969); on moose dung in Canada, Norway and Sweden
(Lundqvist 1972; Kruys etal. 2014).
Notes: We re-examined authentic material (S-F122911) of
Apodospora simulans which was determined by Cain (men-
tioned in the label of material).
Bombardia (Fr.) P. Karst., Bidr. Känn. Finl. Nat. Folk
23: 20 (1873)
Saprobic on wood. Sexual morph: Ascomata perithecial,
immersed to semi-immersed, solitary or gregarious, oval to
subglobose, coriaceous to membranaceous, dark brown to
black, glabrous or surrounded by branched hairs, ostiolate,
with papilla. Paraphyses filiform, branched. Asci 8-spored,
unitunicate, ellipsoidal to cylindrical, pedicellate, rounded or
truncate apex, evanescent. Ascospores hyaline and cylindri-
cal when young, upper cell becoming swollen to fusiform or
oval, brown; lower cell cylindrical or subglobose, hyaline,
smooth-walled, with guttules. Asexual morph: Undeter-
mined (adapted from Fries 1849).
Notes: Fries (1849) introduced Sphaeria subgenus,
Bombardia, based on ‘Sphaeria (Bombardia) fasciculata’,
and Karsten (1873) raised this species to genus level and
established Bombardia. Saccardo (1882) clarified ‘Sphaeria
(Bombardia) fasciculata’ (≡ Sphaeria fasciculata Fr.) and
S. bombarda Batsch. as conspecific, and named Bombar-
dia fasciculata which is a homonym of B. fasciculata Rab.
(Batsch 1786; Rabenhorst 1851). However, the illegitimate
B. fasciculata is still used (Maharachchikumbura etal.
2016; Index Fungorum 2020) and caused some confusion.
Bombardia bombarda (= Sphaeria bombarda Batsch.) was
recommended as the generic type instead of B. fasciculata
(Schröter 1894; Lundqvist 1972), and its molecular data was
provided later (Miller and Huhndorf 2005; Spatafora etal.
2006). In this study, Bombardia is sister to Bombardioidea
in Bombardiaceae (100%ML/1.00PP, Fig.26).
Type species: Bombardia bombarda (Batsch) J. Schröt.,
in Cohn, Krypt.-Fl. Schlesien (Breslau) 3.2(1–2): 302 (1893)
Basionym: Sphaeria bombarda Batsch, Elench. fung.
(Halle): 271 (1786)
Facesoffungi number: FoF 10016; Fig.28
Saprobic on wood. Sexual morph: Ascomata
1–1.5 × 0.4–0.8 mm (x̄ = 1.2 × 0.6 mm, n = 5), per ithe-
cial, immersed to semi-immersed, gregarious, oval to
subglobose, coriaceous, dark brown to black, glabrous,
ostiolate papilla, with 2.5µm wide periphyses. Peridium
75–120µm (x̄ = 95 µm, n = 30) wide, cor iaceous, com-
prising four layers, outer layer composed of amorphous
brown to black material; middle layer composed of hya-
line to brown cells of textura intricata, and dark brown,
strongly tight cells of textura angularis; inner layer com-
posed of pale brown to hyaline cells of textura prismatica.
Paraphyses 2–3µm wide, filiform, septate, evanescent.
Asci (180–)200–260(–310) × 10–15µm (x̄ = 230 × 12µm,
n = 30), 8-spored, unitunicate, ellipsoidal to cylindrical,
pedicellate, apex rounded, with apical globule, evanes-
cent. Ascospores bi-seriate, hyaline and cylindrical when
young, upper cell becoming swollen to fusiform, brown,
(10–)12–15 × 5–10µm (x̄ = 13.5 × 8µm, n = 50), with an api-
cal germ pore, sometimes with a tiny, hyaline, unconspicu-
ous decoration on the apex; lower cell cylindrical, hyaline,
(20–)24–30 × 3.5–6µm (x̄ = 26 × 4.5µm, n = 50), smooth-
walled, guttulate, with a terminal conical appendage at the
base, evanescent. Asexual morph: Undetermined.
Material examined: Italy, Veneto, Treviso, Cansiglio, on
dead trunks of Fagus, October 1874, P.A. Saccardo (S-F750);
Ukraine, Lvivska oblast, Stryi, riverbank near Wierzany, on
stems of Alnus, 28 October 1916, F. Petrak (S-F753).
Known hosts and distribution: On dead, decorticated
wood in Germany (type locality) (Saccardo 1882); on dead
wood in Belgium, Denmark, England, Finland, France, Ger-
many, Hungary, Italy, New Zealand, Poland, Russia, Swe-
den, UK, Ukraine and USA (Lundqvist 1972; Miller and
Huhndorf 2005).
Notes: We re-examined authentic materials of Bombar-
dia bombarda, F750 and F753, collected by Saccardo and
Petrak respectively. The molecular data of B. bombarda were
provided (Miller and Huhndorf 2005; Spatafora etal. 2006),
and in this study, B. bombarda (SMH4821) is sister to Bom-
bardioidea anartia (HHB99-1) (100%ML/1.00PP, Fig.26).
Bombardioidea C. Moreau ex N. Lundq., Symb. bot.
upsal. 20(no. 1): 274 (1972)
Coprophilous. Sexual morph: Ascomata perithecial,
superficial or semi-immersed, gregarious or scattered,
ovoid to subglobose, coriaceous to membranaceous, dark
brown to black, verrucose, ostiolate, with papilla. Peridium
coriaceous, outer layer composed of amorphous brown to
black material; middle layer composed of hyaline to brown
cells of textura intricata, and dark brown, strongly tight
cells of textura angularis; inner layer composed of hyaline
cells of textura prismatica. Paraphyses filiform, septate.
Asci 4–8-spored, unitunicate, ellipsoidal to cylindrical,
pedicellate, usually with apical ring distinct, evanescent.
Ascospores uni- or bi-seriate, ellipsoidal, ovoid or oblong,
hyaline when young, becoming brown when mature, asep-
tate, verrucose, with a germ pore at each end, surrounded by
a gelatinous sheath, guttulate. Asexual morph: Hyphomyce-
tous, phialophora-like. Conidiophores brown to dark brown,
septate, branched, clusters of several phialides, producing
conidia in chains or forming globose masses on the apex.
Conidia numerous, aseptate, hyaline to pale brown, globose
to subglobose (adapted from Niessl 1872; Moreau 1953;
Lundqvist 1972).
Fungal Diversity
1 3
Notes: Sordaria bombardioides was transferred to Bom-
bardioidea as its generic type (Niessl 1872; Moreau 1953;
Lundqvist 1972). Species of this genus are coprophilous usu-
ally reported on leporid dung (Lundqvist 1972; Krug and
Scott 1994). Huhndorf etal. (2004b) sequenced Bombar-
dioidea anartia and in this study, it is sister to Bombardia
bombarda (100%/ML/1.00PP) in Bombardiaceae (Fig.26).
Type species: Bombardioidea bombardioides (Auersw.)
in Lundqvist, Symb. bot. upsal. 20(no. 1): 277 (1972)
Basionym: Sordaria bombardioides Auersw., in Niessl,
Verh. nat. Ver. Brünn 10: 187 (1872)
Facesoffungi number: FoF 10017; Fig.29
Coprophilous. Sexual morph: Ascomata
0.5–1.5 × 0.5–0.8mm (x̄ = 1.2 × 0.6mm, n = 5), perithecial,
superficial, gregarious, oval to subglobose, coriaceous, dark
brown to black, verrucose, ostiolate, with papilla. Peridium
120–160µm (x̄ = 135µm, n = 30) wide, coriaceous, compris-
ing four layers, outer layer composed of amorphous brown to
Fig. 28 Bombardia bombarda:
a, b, d, g–i, m–p (S-F750); c, e,
f, j–l, q–t (S-F753). a Her-
barium material. b, c Ascomata
on host. d, e Ascoma in cross
section. f Ostiole with peri-
physes. g Peridium. h–l Asci.
m–t Ascospores. Scale bars:
b–c = 1mm, d–e = 200µm,
f. h–l = 100µm, g = 50µm,
m–t = 20µm
Fungal Diversity
1 3
black material; middle layer composed of hyaline to brown
cells of textura intricata, and dark brown, strongly tight
cells of textura angularis; inner layer composed of hyaline
cells of textura prismatica. Paraphyses filiform, septate.
Asci (140–)145–160 × 15–25µm (x̄ = 150 × 20 µm, n = 30)
ascospore-bearing part, 8-spored, unitunicate, ellipsoidal to
cylindrical, pedicellate, about 80µm long, apex rounded,
evanescent. Ascospores (20–)22–27(–30) × 10–17 µm
(x̄ = 25 × 12µm, n = 50), uni-seriate, ellipsoidal, hyaline
when young, becoming brown when mature, aseptate, verru-
cose, with a germ pore at each end, surrounded by gelatinous
sheath 1–6µm wide, guttulate. Asexual morph: Hypho-
mycetous. Conidiophores brown to dark brown, becoming
hyaline towards the apex, septate, branched, with clusters of
Fig. 29 Bombardioidea
bombardioides: a, b, e, f, h–j,
n–p (S-F726); c, d, g, k–m (S-
F729); r (redrawn from Krug
and Scott 1994). a Herbarium
material. b, c Ascomata on
host. d Ascoma in cross sec-
tion. e Ostiole with periphyses.
f, g Peridium. h–k Asci. l–q
Ascospores (arrowed: germ
pore). r Conidiophores with
conidia. Notes: Fig j–k. n–p
stained in Congo red. Scale
bars: c = 500µm, d = 200µm,
f. g = 100µm, e. h–k = 50µm,
l–q = 20µm, r = 5µm
Fungal Diversity
1 3
2–7 phialides. Phialides subhyaline to dark brown, subglo-
bose to ampulliform. Conidia numerous, aseptate, hyaline
to pale brown, globose to subglobose (adapted from Krug
and Scott 1994).
Material examined: Czech Republic, Jihomoravsky
Kraj, Brünn, on hare dung, 9 August 1883, G. Niessl von
(S-F726); Poland, Silesia, Karlsbrunn, on hare dung, August
1902, G. Niess von (S-F729).
Known hosts and distribution: On hare dung in Germany
(type locality) (Niessl 1872); on hare and rabbit dung, occa-
sionally on cow, roe-deer, horse, mose, porcupine and sheep
dung in Austria, Bulgaria, Canada, Croatia, Czech Republic,
Denmark, England, Japan, Israel, Germany, Hungary, Neth-
erlands, Poland, Sweden and USA (Lundqvist 1972; Krug
and Scott 1994).
Notes: We re-examined authentic specimens of Bombar-
dioidea bombardioides, F726 and F729, collected by Niessl
and determined by Lundqvist (mentioned in the label of
material). In this study, we provided hand-drawings of the
asexual morph (Fig.29r) from Krug and Scott (1994).
Fimetariella N. Lundq., Bot. Notiser 117: 239 (1964)
Coprophilous. Sexual morph: Ascomata perithecial,
solitary to scattered, immersed to semi-immersed, pyri-
form to subglobose, brown to black, ostiolate. Peridium
comprising membranaceous to coriaceous, pale brown to
brown cells. Paraphyses numerous, filiform, evanescent.
Asci 4–8-spored, unitunicate, cylindrical, pedicellate, apex
rounded. Ascospores uni-seriate, ellipsoidal to oval, asep-
tate, hyaline to brown to dark brown, with a germ pore at
one or each end, surrounded by hyaline, gelatinous sheath.
Asexual morph: Hyphomycetous. Phialides abundant,
hyaline, reduced to collarettes producing conidia. Conidia
ovoid, hyaline, aseptate (adapted from Lundqvist 1964b;
Kruys etal. 2014).
Notes: Lundqvist (1964b) transferred Sordaria rabenhrstii
to Fimetariella as its type species. It is characterized by pyri-
form ascomata and aseptate ascospores surrounded by a
hyaline, gelatinous sheath. Kruys etal. (2014) sequenced F.
rabenhorstii (20410), which was isolated from moose dung
in Sweden and collected by Lundqvist. This strain is basal
to Podospora fabiformis and P. fibrinocaudata in Bombar-
diaceae (100%ML/1.00PP, Fig.26).
Type species: Fimetariella rabenhorstii (Niessl) N. Lundq.,
Bot. Notiser 117: 239 (1964)
Basionym: Sordaria rabenhorstii Niessl, in Rabenhorst,
Fungi europ. exsicc.: no. 1528 (1873)
Facesoffungi number: FoF 10018; Fig.30
Coprophilous. Sexual morph: Ascomata
400–500 × 350–400 µm (x̄ = 430 × 380 µm, n = 10), peri-
thecial, solitary, immersed to semi-immersed, ovoid to sub-
globose, membranaceous, black, ostiolate, with papilla.
Peridium 24–40µm (x̄ = 30 µm, n = 30) wide, outer layer
composed of pale brown to brown cells of textura angula-
ris; inner layer composed of hyaline cells of textura pris-
matica. Paraphyses numerous, filiform, evanescent, broader
towards the base, without constrictions, septate. Asci
(95–)110–128(–132) × 9.5–13µm (x̄ = 120 × 12µm, n = 20),
4-spored, unitunicate, cylindrical, pedicellate, apex rounded
with thickened wall. Ascospores (10–)13–15(–18) × 7.5–10µm
(x̄ = 14 × 8.5µm, n = 40), uni-seriate, ellipsoidal to oval, asep-
tate, hyaline when young, becoming brown to dark brown,
surrounded by a hyaline, gelatinous sheath 1–2.5µm wide.
Asexual morph: Undetermined.
Material examined: Sweden, Uppland, Älvkarleby par.,
SE of the mire Gustavmyrarna (4km SW of Skutskär), in
coniferous forest, on elk dung (Alces alces) in moist cham-
ber, 8 June 1996, N. Lundqvist (S-F66877); Czech Republic,
Jihomoravsky kraj, Brünn, on dung of Caprearum, G. Niessl
(S-F66507, holotype); Czech Republic, Jihomoravsky
kraj, Brünn, on dung of Caprearum, G. Niessl (S-F66508,
isotype).
Known hosts and distribution: On dung of roe and hare in
the Czech Republic (type locality) (Saccardo 1882); on dung
of deer, elk, goat, moose, rabbit, wapiti in Belgium, Canada,
Denmark, Sweden and USA (Krug 1995).
Notes: We re-examined the type materials of Sord-
aria rabenhorstii, which were determined as Fimetariella
rabenhrstii by Lundqvist (mentioned in the label of mate-
rial). It is in good condition.
Ramophialophora M. Calduch, Stchigel, Gené & Guarro,
Stud. Mycol. 50(1): 84 (2004)
Saprobic on wood and/or isolated from soil. Sexual
morph: Undetermined. Asexual morph: Hyphomycetous.
Mycelium superficial to immersed. Hyphae pale brown,
septate, branched. Conidiophores macronematous, mon-
onematous, erect, cylindrical, septate, branched, brown to
dark brown. Conidiogenous cells monophialidic or polyphi-
alidic, terminal or/and lateral. Conidia brown, globose to
oval, aseptate, with a protuberant basal hilum (adapted from
Calduch etal. 2004).
Notes: Ramophialophora is typified by R. vesiculosa
and has macronematous, mononematous conidiophores
and aseptate, brown conidia in slimy masses (Calduch
etal. 2004). The genus was accepted as a member of
Lasiosphaeriaceae based on phylogenetic analysis (Hyde
etal. 2020; Vu etal. 2019; Wijayawardene etal. 2020). In
this study, the strains of Ramophialophora are scattered
in Bombardiaceae, Lasiosphaeriaceae, Neoschizotheci-
aceae and Zygospermellaceae in Sordariales (Fig.26).
However, the generic type R. vesiculosa (CBS 110629) is
sister to Bombardia and Bombardioidea (100%ML/1.00PP,
Fig.26). Therefore, we recommend transferring this genus
to Bombardiaceae.
Fungal Diversity
1 3
Type species: Ramophialophora vesiculosa M. Calduch,
Stchigel, Gené & Guarro, in Calduch, Gené, Cano, Stchigel,
Cano & Guarro, Stud. Mycol. 50(1): 84 (2004)
Facesoffungi number: FoF 10045; Fig.31
Saprobic on wood and/or isolated from soil. Sexual
morph: Undetermined. Asexual morph: Hyphomycetous.
Colonies on sterilized wood, hairy, brown. Mycelium super-
ficial to immersed on substrate. Hyphae 1.5–2.5µm wide,
pale brown, septate, branched. Conidiophores 2–6.5µm
wide, macronematous, mononematous, erect, cylindrical,
septate, branched, pale brown to dark brown, becoming paler
towards the apex. Conidiogenous cells 5.5–9 × 2.5–4 µm
(x̄ = 7 × 3.5µm, n = 30), monophialidic or polyphialidic,
terminal and lateral, lageniform, smooth-walled, with con-
spicuous collarettes, 1.5–2.5µm wide. Conidia 2–3µm
(x̄ = 2.5µm, n = 30) diam., aggregated, pale brown to brown,
Fig. 30 Fimetariella rabenhorstii: a, b, d, f, m, n, q, r (S-F66507,
holotype); c, e, g–l, o, p (S-F66508, isotype). a Herbarium material.
b, c Ascomata on host. d Ascoma in cross section. e Ostiole with hya-
line periphyses. f, g Peridium. h–j Asci. k–r Ascospores. Scale bars:
b. c = 500µm, d = 200µm, e–j = 50µm, k–r = 10µm
Fungal Diversity
1 3
globose to ellipsoidal, aseptate, smooth-walled, with a trun-
cate and protuberant base.
Material examined: Spain, Asturias, Muniellos Integral
Biological Reserve, in soil (isolated on sterilized wood), 26
June 1999, M. Calduch and A.M. Stchigel (IMI-389151,
holotype).
Known hosts and distribution: On soil in Spain (type
locality) (Calduch etal. 2004).
Lasiosphaeriaceae Nannf., Nova Acta R. Soc. Scient.
upsal., Ser. 4 8(no. 2): 50 (1932)
Coprophilous or saprobic on wood or isolated from soil.
Sexual morph: Ascomata perithecial or cleistothecial, soli-
tary to scattered or gregarious, immersed to semi-immersed
to superficial, globose to subglobose, dark brown to black.
Paraphyses numerous, filiform to cylindrical, septate, eva-
nescent. Asci 4–8-spored, unitunicate, cylindrical to clavate,
pedicellate. Ascospores uni-seriate or bi-seriate, ellipsoi-
dal to oval or irregular, aseptate to multi-septate, hyaline
to brown to dark brown, verrucose or smooth-walled, with
a gelatinous sheath or irregular appendage(s). Asexual
morph: Hyphomycetous. Conidiophores arising from aerial
hyphae, erect, inflated at the base. Conidia small, oval to
globose, ovate forming globose masses on the apex of the
phialides (adapted from Nannfeldt 1932; Lundqvist 1972).
Type genus: Lasiosphaeria Ces. & De Not. 1863
Notes: Lasiosphaeriaceae is typified by Lasiosphaeria
and has black ascomata, cylindrical to clavate asci and
varied ascospores (Cesati and de Notaris 1863; Nannfeldt
1932; Lundqvist 1972; Wang etal. 2019a, b; Marin-Felix
etal. 2020). Species in Lasiosphaeriaceae were reported
to be similar to taxa in Nitschkiaceae, Sordariaceae and
Tripterosporaceae based on their ascospores with vari-
ous ornamentations and appendages, such as Bombardia,
Fig. 31 Ramophialophora vesiculosa (IMI-389151, holotype). a Herbaium material label. b Herbarium material. c Mycelium on host. d Myce-
lium. e–i Conidiophores with conidia. j, k Conidia. Scale bars: d = 100µm, e. f, h. i = 5µm, g = 10µm, j. k = 2µm
Fungal Diversity
1 3
Lasiosphaeria (Munk 1957; Carroll and Munk 1964; Lun-
dqvist 1972; Barr 1990; Kruys etal. 2014). The traditional
classification of Lasiosphaeriaceae is based on morphol-
ogy; obviously, it is inconsistent with molecular phylogeny
(Miller and Huhndorf 2004a, b; 2005; Wang etal. 2019a,
b; Marin-Felix etal. 2020). Huhndorf etal. (2004b) con-
firmed that Lasiosphaeriaceae is polyphyletic based on
LSU sequence data. Subsequently, many polyphyletic gen-
era in this family were repositioned based on the sequence
data (Cai etal. 2006a; Chang etal. 2010; Kruys etal. 2014;
Wang etal. 2019a, b; Marin-Felix etal. 2020). Wang etal.
(2019a, b) redefined three genera, Cladorrhinum, Podos-
pora and Triangularia, and established Podosporaceae
based on phylogenetic analysis. Wijayawardene etal.
(2020) listed 32 genera in Lasiosphaeriaceae. Marin-Felix
etal. (2020) re-classified Cercophora, Diplogelasinos-
pora, Immersiella, Jugulospora, Rinaldiella, Schizothe-
cium, Zygopleurage and some species of Apiosordaria and
Podospora to establish Diplogelasinosporaceae, Navicu-
lisporaceae and Schizotheciaceae in Sordariales based on
morphology and phylogenetic analyses.
In this study, based on phylogenetic analyses, we divide
the remaining Lasiosphaeriaceae taxa into five clades
(Fig.26): (i) Bombardiaceae clade: This is a sister clade
of Sordariaceae (74%ML). Apodospora, Bombardia, Bom-
bardioidea, Fimetariella and some species of Cercophora,
Podospora, Ramophialophora and Zopfiella clustered in this
clade (75%ML); (ii) Lasiosphaeriaceae clade: This clade is
regarded as Lasiosphaeriaceae sensu stricto (Marin-Felix
etal. 2020) because several typical Lasiosphaeria species
with molecular data are included in this clade; however, the
ex-type strain of L. ovina has not been sequenced. This clade
is sister to Naviculisporaceae (69%ML). It clusters Anopo-
dium, Bellojisia, Corylomyces, Lasiosphaeria, Mammaria,
Zopfiella and some species of Cercophora, Podospora and
Ramophialophora in 92%ML support; (iii) Lasiosphaeri-
daceae clade: This is an orphan clade in Sordariales, which
clusters Lasiosphaeris species only (100%ML/1.00PP);
(iv) Strattoniaceae clade: This is an unstable clade which
includes Strattonia species and is sister to Diplogelasino-
sporaceae with poor support; (v) Zygospermellaceae clade:
This clade contains Episternus onthosphagi, Zygospermella
insignis and Ramophialophora petraea (83%ML/0.99PP),
and is an orphan clade in Sordariales (83%ML/1.00PP).
In addition, the generic type of Apodus is nested in
Neoschizotheciaceae and, the type species of Ramophial-
ophora gathers in Bombardiaceae in phylogenetic analysis
(Fig.26). The polyphyletic Arnium, which has no molecular
data of their type species, are placed in Sordariales genera
incertae sedis. Given the diversity of the morphology of
lasiosphaeriaceous taxa and locating each genus more accu-
rately, we put the genera (Biconiosporella, Camptosphaeria,
Diffractella, Emblemospora, Eosphaeria, Periamphispora,
Tripterosporella) without molecular data in Sordariales gen-
era incertae sedis pending further studies.
Thaxteria is typified by T. didyma, which was syn-
onymized under Bertia didyma (Saccardo 1891; Mugambi
and Huhndorf 2010). Apiosordaria is the synonym of Trian-
gularia based on its available molecular data of type species
(Wang etal. 2019a, b). Therefore, they will not be discussed
in this study.
Marin-Felix etal. (2020) redefined Lasiosphaeriaceae and
proposed that Lasiosphaeriaceae sensu stricto may divided
into two clades; Lasiosphaeria clade and the clade includes
the generic type of Anopodium, Bellojisia, Corylomyces,
Zopfiella and several other species Cercophora sulphurella,
C. sparsa, Podospora didyma and Ramophialophora humi-
cola (clade I, Fig.26). In this study, we added the molecular
data which was derived from Vu etal. (2019) of Mammaria
and found that this genus is basal to Lasiosphaeria clade and
clade I in Lasiosphaeriaceae (86%ML, Fig.26). Therefore,
six genera are accommodated in Lasiosphaeriaceae based
on phylogenetic analysis.
This family has high morphological variability. Ramo-
phialophora humicola is a phialophora-like hyphomycetes
(Madrid etal. 2010). All sexual species have perithecial
ascomata, except for Zopfiella tabulata, which has cleis-
tothecial ascomata; Anopodium ampullaceum, Cercophora
sulphurella, C. sparsa, Mammaria echinnobotryoides,
Podospora didyma and Zopfiella tabulata have cylin-
drical ascospores with swollen apical cell and terminal
appendage(s) present or absent, whereas Bellojisia rhyn-
chostoma and Corylomyces seleonsporus have allantoid
to reniform ascospores (Saccardo 1878, 1906; Lundqvist
1964a, b, 1972; Guarro etal. 1991; Stchigel etal. 2006;
Réblová 2008). However, more fresh collections and phy-
logenetic data are requested to determine the affinities and
taxonomic placement for the members in this clade.
Anopodium N. Lundq., Bot. Notiser 117: 356 (1964)
Coprophilous. Sexual morph: Ascomata perithecial,
scattered, semi-immersed, or erumpent through the bark
of host, or superficial, globose, pale brown to dark brown,
ostiolate, with black papilla, surrounded by setae. Peridium
membranaceous, composed of hyaline to brown cells of
textura angularis to textura prismatica. Asci 8-spored, unitu-
nicate, ellipsoidal, pedicellate, blunt at the apex. Ascospores
subglobose to oval, collapsing when dry, aseptate, hyaline
when young, becoming dark brown, smooth-walled, with an
apical germ pore. Asexual morph: Undetermined (adapted
from Lundqvist 1964a).
Notes: Anopodium was introduced by Lundqvist (1964a)
with two species A. ampullaceum (type) and A. epile. This
coprophilous genus is characterized by pale brown asco-
mata with black necks and ascospores with a pedicel at
the end (Lundqvist 1964a). Anopodium epile differs from
Fungal Diversity
1 3
A. ampullaceum in having glabrous ascomata (Lundqvist
1964a). Kruys etal. (2014) sequenced A. ampullaceum. In
this study, Fig.26 shows Anopodium nests in Lasiospha-
eriaceae clade I.
Type species: Anopodium ampullaceum N. Lundq., Bot.
Notiser 117: 356 (1964)
Facesoffungi number: FoF 10019; Fig.32
Coprophilous. Sexual morph: Ascomata
400–550 × 275–400 µm (x̄ = 480 × 335 µm, n = 10),
perithecial, scattered, semi-immersed, or erumpent
through bark of host, or superficial, globose, pale brown
to dark brown, ostiolate, with black papilla, surrounded
by setae 1.5–4.5 wide. Necks 180–200 × 100–135 µm
(x̄ = 185 × 120 µm, n = 5), membranaceous, central,
dark brown to black, smooth-walled, with hyaline peri-
physes. Peridium 20–55µm (x̄ = 40 µm, n = 20) wide,
membranaceous, comprising two layers, outer layer
composed of brown cells of textura angularis; inner
layer composed of hyaline cells of textura prismatica.
Fig. 32 Anopodium ampullaceum: a, b, d, h, j–n (F 47,944); c,
e–g, i (F 66,862). a Herbarium material. b, c Ascomata on host. d,
e Ascoma in cross section. f Black neck in cross section. g Peridium.
h–j Asci. k–n Ascospores (arrows show germ pores). Scale bars:
b = 500µm, c–e = 200µm, f–g = 100µm, h–j = 50µm, k–n = 20µm
Fungal Diversity
1 3
Paraphyses 4–5µm wide, numerous, broad, evanescent.
Asci (135–)150–170(–200) × 35–45µm (x̄ = 160 × 40µm,
n = 30), 8-spored, unitunicate, ellipsoidal, pedicellate,
apex blunt, apical ring absent, evanescent. Ascospores
(25–)28–32(–35) × 15–25 µm (x̄ = 30 × 20 µm, n = 50)
for the mature swollen cells, bi-seriate, subglobose to
oval, collapsing when dry, aseptate, hyaline, swollen cell
towards the apex of ascus when young, becoming dark
brown, partly ascospores of their swollen cell toward
the apex; partly toward the base when mature, smooth-
walled, with an apical germ pore and an evanescent pedicel
(10–)18–25 × 1.5–3.5µm (x̄ = 20 × 2.5µm, n = 20) at the
other end. Asexual morph: Undetermined.
Material examined: Sweden, Jämtland, Mårdsunds-
bodarna, Hallen par., in subalpine birch forest, on hare dung
(Lepus timidus) in moist chamber, Stockholm, 11 July 1988,
N. Lundqvist (S-F47944); UK, Midlothian, Edinburgh,
Hillend, on rabbit dung (Oryctolagus cuniculus) in moist
chamber, 26 December 1996, M.J. Richardson (S-F66862).
Known hosts and distribution: On fresh hare dung in
Sweden (type locality) (Lundqvist 1964a); on rabbit dung in
Belgium, Norway, Sweden and UK (Lundqvist 1972; Kruys
etal. 2014).
Notes: Lundqvist (1964a) discovered Anopodium ampul-
laceum on hare dung in Sweden, and it was reported in
the UK (Kruys etal. 2014). In this study, we re-examined
authentic material collected by Lundqvist in Sweden and a
sample collected in the UK. We found that most of the swol-
len ascospores cells in the same ascus from S-F47944 were
towards the ascus apex when they were young, becoming
partly toward the ascus apex and partly toward the base.
Kruys etal. (2014) sequenced A. ampullaceum (MJR 40/07
and E00218015) and found that these strains are related to
Bellojisia, Corylomyces and Lasiosphaeria. In this study, A.
ampullaceum (MJR 40/07) is nested in Lasiosphaeriaceae
clade I (66%ML, Fig.26).
Bellojisia Réblová, Mycologia 100(6): 897 (2008)
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, solitary to gregarious, semi-immersed to superficial,
carbonaceous, pyriform to subglobose, tomentose, brown
to black, ostiolate, with necks, periphysate. Peridium com-
posed of brown to hyaline cells of textura prismatica to
textura intricata to textura angularis. Paraphyses septate.
Asci 8-spored, unitunicate, cylindrical, pedicellate, with J-
apical ring. Ascospores 0–1-septate, navicular to reniform
to falciform, dark brown, with a large guttule in each cell
and an apical germ pore. Asexual morph: Undetermined
(adapted from Réblová 2008).
Notes: The monotypic Bellojisia was established based
on Jobellisia rhynchostoma in phylogenetic analysis and
its unique reniform ascospores different from Jobellisia
(Réblová 2008). In this study, the strain of B. rhynchostoma
is basal to Podospora didyma and Zopfiella tabulata in Lasi-
osphaeriaceae clade I (100%ML/0.95PP, Fig.26).
Type species: Bellojisia rhynchostoma (Höhn.) Réblová,
Mycologia 100(6): 897 (2008)
Facesoffungi number: FoF 10123; Fig.33e–f
Saprobic on wood. Sexual morph: Ascomata perithecial,
solitary to gregarious, semi-immersed to superficial, carbo-
naceous, pyriform to subglobose, tomentose, brown to black,
ostiolate, with necks, periphysate. Peridium comprising two
layers, outer layer composed of brown cells of textura pris-
matica to textura intricata, inner layer composed of hyaline
to brown cells of textura angularis. Paraphyses septate. Asci
8-spored, unitunicate, cylindrical, pedicellate, with J- apical
ring. Ascospores 0–1-septate, navicular to reniform to fal-
ciform, dark brown, with a large guttule in each cell and an
apical germ pore. Asexual morph: Undetermined (adapted
from Réblová 2008).
Known hosts and distribution: In pericarps of Juglans
regia in the Austria (type locality); on decorticated wood of
Robinia pseudoacacia in the France (Réblová 2008).
Notes: The reniform ascospores of Bellojisia rhynchos-
toma are similar to Corylomyces selenosporus and they
nested in Lasiosphaeriaceae clade I in phylogenetic analysis
(Fig.26).
Corylomyces Stchigel, M. Calduch & Guarro, Mycol.
Res. 110(11): 1362 (2006)
Saprobic on wood. Sexual morph: Ascomata perithecial,
gregarious, superficial, pyriform to subglobose, olivaceous-
yellow to olive, tomentose, ostiolate, with long necks, sur-
rounded by hairs. Asci 8-spored, unitunicate, cylindrical,
pedicellate, evanescent. Ascospores uni-seriate, reniform,
0–1-septate, hyaline to dark brown, with an apical, umbonate
germ pore. Asexual morph: Undetermined (adapted from
Stchigel etal. 2006).
Notes: The monotypic Corylomyces was reported on
hazelnut and is placed in Lasiosphaeriaceae based on the
LSU sequence data (Stchigel etal. 2006). Marin-Felix etal.
(2020) recognized that Corylomyces clustered with Lasio-
sphaeriaceae s. str. clade in LSU-ITS-RPB2-TUB sequence
analysis. In this study, the ex-type of C. selenosporus (CBS
113930) is sister to Cercophora sparsa in Lasiosphaeriaceae
clade I (89%ML, Fig.26).
Type species: Corylomyces selenosporus Stchigel, M. Cal-
duch & Guarro [as 'selenospora'], in Stchigel, Cano, Miller,
Calduch & Guarro, Mycol. Res. 110(11): 1362 (2006)
Facesoffungi number: FoF 10124; Fig.34a
Saprobic on wood. Sexual morph: Ascomata perithecial,
gregarious, superficial, pyriform to subglobose, olivaceous-
yellow to olive, tomentose, ostiolate, with long necks, sur-
rounded by hairs. Asci 8-spored, unitunicate, cylindrical,
pedicellate, evanescent. Ascospores uni-seriate, reniform
Fungal Diversity
1 3
to lunate, 0–1-septate, hyaline to dark brown, with an api-
cal, umbonate germ pore. Asexual morph: Undetermined
(adapted from Stchigel etal. 2006).
Known hosts and distribution: On hazelnut (Corylus avel-
lana) decomposing on soil in France (type locality) (Stchigel
etal. 2006).
Fig. 33 Podospora fimicola: a, b (S-F26041, lectotype); c, d
(redrawn from Bell and Mahoney 1997; Doveri 2008). a Herbarium
material. b Ascoma on host. c Ascus. d Ascospore; Bellojisia rhyn-
chostoma: e, f (redrawn from Réblová 2008). e Ascus and paraphy-
sis. f Ascospores; Apiosordaria tetraspora: g, h (redrawn from Krug
et al. 1983). g Ascospore. h Conidiophore and conidia; Triangu-
laria verruculosa – as Apiosordaria verruculosa: i, j (redrawn from
Jensen 1912). i Ascus. j Ascospore. Scale bars: c. d = 100 µm, e. h.
i = 20µm, f. g. j = 10µm
Fig. 34 Corylomyces selenosporus: a (redrawn from Stchigel etal. 2006). a Ascospores; Diplogelasinospora princeps: b (redrawn from Cain
1961). b Ascospores; Zopfiella tabulata: c (redrawn from Winter 1884). c Ascospores. Scale bars: a–c = 10µm
Fungal Diversity
1 3
Notes: Bellojisia rhynchostoma and Corylomyces seleno-
sporus have long necks, tomentose ascomata and 0–1-septate
ascospores; whereas the ascospores of the former are lunate
and the latter are navicular (Stchigel etal. 2006; Réblová
2008). Cercophora sparsa has cylindical ascospores, which
is different from Cor. selenosporus has navicular ascospores
(Hilber and Hilber 1979; Stchigel etal. 2006).
Lasiosphaeria Ces. & De Not., Comm. Soc. crittog. Ital.
1(fasc. 4): 229 (1863)
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, solitary or gregarious, superficial to semi-immersed,
globose to subglobose, brown, surrounded by white furfura-
ceous cover, ostiolate, with distinct black papilla. Peridium
comprising membranaceous to coriaceous, brown to hya-
line cells. Paraphyses filiform. Asci 8-spored, unitunicate,
cylindrical, pedicellate. Ascospores cylindrical to ellipsoi-
dal, sometimes slight curved, hyaline to brown, aseptate to
multi-septate, guttulate, smooth-walled. Asexual morph:
Hyphomycetous, phialophora-like. Conidiogenous cells phi-
alides, monophialidic, cylindrical to oblong, hyaline, with
collarette or collarette absent. Conidia oval to pyriform, hya-
line, truncate at the base (adapted from Cesati and de Notaris
1863; Huhndorf etal. 2004b; Miller and Huhndorf 2004a;
Miller etal. 2014).
Notes: Lasiosphaeria as the type genus of Lasiospha-
eriaceae is characterized by globose ascomata, cylindrical
asci and cylindrical to ellipsoidal ascospores (Cesati and de
Notaris 1863; Huhndorf etal. 2004b; Miller and Huhndorf
2004a; Miller etal. 2014). Strains of Lasiosphaeria spe-
cies are close to Anopodium, Bellojisia, Corylomyces and
Zopfiella in phylogenetic analysis (Maharachchikumbura
etal. 2016; Hyde etal. 2020; Marin-Felix etal. 2020). In
this study, the Lasiosphaeria strains is sister to clade I com-
prises Anopodium, Bellojisia, Corylomyces and Zopfiella
(86%ML), and is related to Mammaria (Fig.26).
Type species: Lasiosphaeria ovina (Pers.) Ces. & De Not.,
Comm. Soc. crittog. Ital. 1(fasc. 4): 229 (1863)
Basionym: Sphaeria ovina Pers., Syn. meth. fung. (Göt-
tingen) 1: 71 (1801)
Facesoffungi number: FoF 01147; Fig.35
Saprobic on wood. Sexual morph: Ascomata
360–450 × 340–370µm (x̄ = 400 × 345µm, n = 10), perithe-
cial, solitary or gregarious, superficial to semi-immersed,
globose to subglobose, brown, membranaceous, surrounded
by white furfuraceous cover, ostiolate, with distinct black
papilla. Peridium comprising three layers, outer layer com-
posed of white cells of textura intricata, 23–48µm wide;
middle layer coriaceous, composed of brown cells of textura
angularis, 23–35µm wide; inner layer membranaceous,
composed of hyaline cells of textura prismatica, 6–16µm
wide. Paraphyses 1–2µm wide, numerous, filiform. Asci
(80–)95–110(–120) × 8–12µm (x̄ = 100 × 10µm, n = 30),
8-spored, unitunicate, cylindrical, pedicellate, apex rounded
with subapical globule, apical ring distinct. Ascospores
(25–)28–34(–45) × 2–5 µm (x̄ = 30 × 3 µm, n = 50), bi-
seriate, slight curved, cylindrical to geniculate, hyaline and
aseptate when young, becoming pale brown and multi-sep-
tate when mature, with a large guttule in each cell, smooth-
walled. Asexual morph: Undetermined.
Material examined: Sweden, Småland, Högsby par., Mas-
semåla, on decayed branch of apple tree in old pasture, 3
October 1992, N. Lundqvist (S-F123316); Sweden, Upp-
land, Uppsala, “Predikstolen” N of Lurbo (5km SSW of
Uppsala), on decayed wood in deciduous forest, 7 September
1994, N. Lundqvist (S-F123318); Sweden, Västergötland,
Säter par., Ăngen (2km WNW of Stöpen), on rotten birch
log in deciduous forest, 13 September 2001, N. Lundqvist
(S-F123375).
Known hosts and distribution: On rotten trunk in Den-
mark, England, Germany (type locality), Sweden, Turkey
and USA (Persoon 1801; Miller and Huhndorf 2004b; Dül-
ger and Akata 2016; Vu etal. 2019).
Notes: Several Lasiosphaeria ovina were sequenced, but
the ex-type strain is lacking (Miller and Huhndorf 2004b;
Fernández etal. 2006; Raja etal. 2011; Vu etal. 2019). In
this study, strains of L. ovina (CBS 126299 and SMH 1538)
are nested in Lasiosphaeria clade (Fig.26). We could not
obtain the type material. Therefore, we re-examined authen-
tic specimens collected by Lundqvist.
Lasiosphaeria rehmiana Henn., Verh. bot. Ver. Prov.
Brandenb. 40: 135 (1898) [1899]
Facesoffungi number: FoF 10020; Fig.36
Saprobic on wood. Sexual morph: Ascomata
220–290 × 240–270µm (x̄ = 240 × 255µm, n = 10), per i-
thecial, gregarious, superficial, globose to subglobose,
brown, membranaceous, surrounded by white furfura-
ceous cover, ostiolate, with distinct black papilla. Setae
3–10µm wide, brown to dark brown, aseptate, swollen at
the base, tapering, with a swollen, hyaline apex. Peridium
comprising three layers, outer layer composed of white
cells of textura intricata, 10–24µm wide; middle layer
coriaceous, composed of brown cells of textura angula-
ris, 10–24µm wide; inner layer membranaceous, com-
posed of hyaline cells of textura prismatica, 6–14µm wide.
Paraphyses 2–4µm wide, numerous, septate, cylindrical.
Asci (100–)115–120(–150) × 10–15µm (x̄ = 118 × 13µm,
n = 30), 8-spored, unitunicate, cylindrical, pedicellate, apex
rounded. Ascospores (20–)24–30(–35) × (3–)4–6(–7) µm
(x̄ = 27 × 5µm, n = 50), bi-seriate, slight curved, ellipsoi-
dal, 3–5-septate, hyaline when young, becoming brown and
swollen at the middle cells when mature, slightly constricted
at septum, guttulate, smooth-walled. Asexual morph:
Undetermined.
Fungal Diversity
1 3
Material examined: Germany, Berlin, Hoard. Bot. Berol.,
Warmhaus, on an imported wood from Cameroon, on
decayed wood, May 1891, P. Hennings (S-F6328, holotype).
Known hosts and distribution: On dead wood in Germany
(type locality) (Hennings 1898).
Notes: Lasiosphaeria rehmiana was introduced as a spe-
cies closely related to Chaetosphaeria in having ellipsoidal,
septate ascospores (Hennings 1898). Réblová and Huhndorf
subsequently re-examined the type material, Lasiospha-
eria rehmiana (F6328) and considered it as Melanochaeta
Fig. 35 Lasiosphaeria ovina: a, b, e, o (S-F123316); h, j–k, n, p
(S-F123318); c, d, f, g, i, l, m, q, r (S-F123375). a Material. b, c
Ascomata on host. d Black papilla of ascoma. e Ascoma in cross sec-
tion. f Peridium. g Paraphyses. h–k Asci. l–r Ascospores. Scale bars:
b–c = 500µm, d = 200µm, e = 100µm, f. h–k = 20µm, g, l–r = 10µm
Fungal Diversity
1 3
hemipsila (Chaetosphaeriaceae) (mentioned in the label of
material).
Mammaria Ces. ex Rabenh., Bot. Ztg. 12: 190 (1854)
Saprobic on wood or leaves. Sexual morph: Ascomata
perithecial, gregarious, superficial, globose to subglobose,
black, ostiolate. Asci 8-spored, unitunicate, cylindrical.
Ascospores bi-seriate, cylindrical to geniculate, hyaline
to brown, with a germ pore at each end. Asexual morph:
Hyphomycetous. Mycelium hyaline to pale brown, branched.
Conidiophores mononematous, erect, hyaline to pale brown,
branched. Conidia globose, hyaline to pale brown, smooth-
walled (adapted from Cesati 1854; del Valle Catania etal.
2011).
Notes: Mammaria was introduced as a hyphomycetous
genus and is typified by M. echinobotryoides (Cesati 1854).
Saccardo (1886) transferred this species to Trichosporum
(Dothideomycetes). Vu etal. (2019) sequenced M. ech-
inobotryoides, and in this study, M. echinobotryoides is
branched out of Lasiosphaeria and clade I in Lasiospha-
eriaceae (86%ML, Fig.26). Pseudocercophora ingoldii and
Cercophora solaris were reported as the sexual morph of
M. echinobotryoides (Subramanian and Sekar 1986; del
Valle Catania etal. 2011), and Réblová etal. (2016) recom-
mended using the old Mammaria over Pseudocercophora.
In this study, Mammaria echinobotryoides and C. sola-
ris clustered (100%ML/1.00PP) in Lasiosphaeriaceae
(Fig.26).
Fig. 36 Lasiosphaeria
rehmiana: (S-F6328, holo-
type). a Herbarium material.
b. Ascomata. c. Ascoma in
cross section. d. Peridium.
e. Seta. f. Paraphyses. g, h.
Asci. i–l. Ascospores. Scale
bars: b = 500µm, c = 100µm,
d–h = 50µm, i–l = 20µm
Fungal Diversity
1 3
Type species: Mammaria echinobotryoides Ces., in Raben-
horst, Klotzschii Herb. Viv. Mycol. 19: no. 1859 (1854)
Synonym: Cercophora solaris (Cooke & Ellis) R. Hilber
& O. Hilber, Z. Mykol. 45(2): 221 (1979)
Facesoffungi number: FoF 10125; Fig.37a–c
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, gregarious, superficial, globose to subglobose, black,
ostiolate, with papilla. Asci 8-spored, unitunicate, cylindri-
cal. Ascospores bi-seriate, aseptate, cylindrical to geniculate,
hyaline, becoming 0–1-septate, upper cell triangular, brown;
lower cell cylindrical, hyaline, with a germ pore at each end.
Asexual morph: Hyphomycetous. Mycelium hyaline to pale
brown, branched. Conidiophores mononematous, erect, hya-
line to pale brown, branched. Chlamydospores oval to ellip-
soid, pale brown to brown, slightly apiculate, flattened basal,
with a longitudinal germ slit. Conidia globose, hyaline to
pale brown, smooth-walled (adapted from Hennebert 1968;
del Valle Catania etal. 2011).
Known hosts and distribution: On hollow trunk in Bel-
gium, Canada, Italy (type locality), Singapore and USA (Sac-
cardo 1886; Subramanian and Sekar 1986; Vu etal. 2019).
Notes: Mammaria echinobotryoides was found as a
hyphomycetous species (Cesati 1854). Later Cercophora
solaris was introduced as the sexual morph of M. echino-
botryoides based on similar cultural characteristics (Sub-
ramanian and Sekar 1986, del Valle Catania etal. 2011).
Cercophora solaris has cylindrical to geniculate ascospores
similar to Lasiosphaeria; phylogenetically, the strains of
Cercophora solaris and M. echinobotryoides clustered
(100%ML/1.00PP, Fig.26). Thus, we recommend to transfer
C. solaris as M. echinobotryoides.
Zopfiella G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig)
1.2: 56 (1884).
Saprobic on wood or coprophilous. Sexual morph: Asco-
mata cleistothecial, scattered, superficial, globose to subglo-
bose, brown, surrounded by septate, olive to yellow-green
hairs. Peridium membranaceous, comprising brown to oli-
vaceous brown cells of textura angularis. Asci 4–8-spored,
unitunicate, cylindrical to clavate, evanescent. Ascospores
bi-seriate, upper cell(s) ellipsoidal to fusiform, aseptate
to multi-septate, pale brown to dark brown, with an apical
Fig. 37 Mammaria echino-
botryoides: a–c (redrawn from
Hennebert 1968; del Valle
Catania etal. 2011). a, b
Conidiophore, Chlamydospores
and conidia. c Ascospores;
Cladorrhinum foecundissi-
mum: d (redrawn from Hyde
and Goh 1999). d Conidiophore
and conidia; Triangularia bam-
busae: e (redrawn from Guarro
and Cano 1988). e Ascospores;
Tripterosporella coprophila: f,
g (redrawn from Subramanian
and Lodha 1968). f Ascospores.
g Ascus; Rinaldiella pentago-
nospora: h, i (redrawn from
Crous etal. 2014). h Ascus.
i Ascospores. Scale bars:
f–h = 20µm, a–e. i = 10µm
Fungal Diversity
1 3
germ pore and a truncate base; lower cell hyaline, cylindri-
cal, gelatinous, evanescent. Asexual morph: Hyphomycet-
ous, humicola-like. Conidia globose, hyaline to pale brown
(adapted from Winter 1884; Guarro etal. 1991).
Notes: Zopfiella is characterized by cleistothecial asco-
mata and ascospores comprise ellipsoidal, brown upper
cell(s) and a cylindrical, hyaline pedicel (Winter 1884;
Guarro etal. 1991). In subsequent phylogenetic analy-
ses Zopfiella species scattered in Naviculisporaceae,
Podosporaceae and Neoschizotheciaceae (Wang etal. 2019a,
b; Marin-Felix etal. 2020; this study, Fig.26). However, the
generic type, Z. tabulata is sister to Podospora didyma in
Lasiosphaeriaceae clade I (100%ML/0.95PP, Fig.26).
Type species: Zopfiella tabulata (Zopf) G. Winter, Rabenh.
Krypt.-Fl., Edn 2 (Leipzig) 1.2: 56 (1884)
Basionym: Cephalotheca tabulata Zopf, Sber. Gesells-
chaft naturf. Freunde Berlin 2: 34 (1880)
Facesoffungi number: FoF 10126; Fig.34c
Coprophilous. Sexual morph: Ascomata cleistothecial,
scattered, superficial, globose to subglobose, brown, sur-
rounded by septate, olive to yellow-green hairs. Peridium
membranaceous, comprising brown to olivaceous brown
cells of textura angularis. Asci 4-spored, unitunicate, cylin-
drical to clavate, evanescent. Ascospores bi-seriate, ellipsoi-
dal, hyaline when young, becoming upper cell(s) ellipsoidal
to fusiform, 0–1-septate, pale brown to dark brown, with an
apical germ pore and a truncate base; lower cell hyaline,
cylindrical, gelatinous, evanescent. Asexual morph: Unde-
termined (adapted from Winter 1884; Guarro etal. 1991).
Known hosts and distribution: On dung of hare, porcu-
pine, rabbit and sheep in Germany (type locality) (Winter
1884; Cai etal. 2005).
Notes: Zopfiella tabulata was isolated from dung of hare,
rabbit and sheep (Winter 1884; Guarro etal. 1991). Subse-
quently, Cai etal. (2005) sequenced Zopfiella tabulata (CBS
230.78) collected from porcupine dung. This strain is sister
to Podospora didyma (CBS 232.78) in phylogenetic analysis,
whereas the latter has perithecial ascomata and ascospores
with terminal appendages (Mirza and Cain 1969).
Lasiosphaeridaceae S.K. Huang, Maharachch. & K.D.
Hyde, fam. nov.
Facesoffungi number: FoF 10142; Index Fungorum num-
ber: IF558345
Etymology: Named for Lasiosphaeris, the type genus of
this family.
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, solitary, superficial to semi-immersed, subglobose to
obpyriform, black, ostiolate, tuberculate or/and surrounded
by brown, septate setae or hairs. Asci 8-spored, unitunicate,
cylindrical, with J- apical ring and a subapical globule.
Ascospores cylindrical to geniculate or sigmoid, aseptate to
multi-septate, ends rounded, hyaline to pale brown, becom-
ing brown to dark brown, ellipsoidal to broadly fusiform at
apical cell(s), sometimes with terminal, short, conical gelati-
nous appendage at each end. Asexual morph: Hyphomy-
cetous, phialophora-like. Conidia oval to globose, hyaline
(adapted from Tode 1791; Hilber and Hilber 1979; Miller
and Huhndorf 2004a).
Type genus: Lasiosphaeris Clem. 1909
Notes: Lasiosphaeris is similar to Lasiosphaeria and
Immersiella in having geniculate or sigmoid ascospores,
but Lasiosphaeria has white furfuraceous tissue covering on
the surface of the ascomata and Immersiella has immersed
ascomata, whereas Lasiosphaeris has superficial, hairy
ascomata (Tode 1791; Clements 1909; Miller and Huhn-
dorf 2004a; Kruys etal. 2014). However, these three gen-
era were subsequently found to be far apart in phylogenetic
analysis (Miller and Huhndorf 2004a; Kruys etal. 2014;
Marin-Felix etal. 2020; this study). Miller and Huhndorf
(2004a) regarded Lasiosphaeris species as an independent
clade in Lasiosphaeriaceae based on LSU sequence data.
Marin-Felix etal. (2020) showed that Lasiosphaeris is sister
to Zygospermella based on multi-gene analysis. In this study,
Cercophora arenicola, Lasiosphaeris hirsuta and L. hispida
form a clade (100%ML/1.00PP) which is nested in Sord-
ariales as an orphan clade (72%ML, Fig.26). Furthermore,
Cercophora arenicola is similar to Lasiosphaeris which has
hairy ascomata, cylindrical to clavate asci with a subapical
globule. Still, ascospores have swollen apical cells with a
terminal appendage at each end (Hilber and Hilber 1979).
Therefore, we accepted C. arenicola as Lasiosphaeris areni-
cola (see the combination list below) and delimited the new
family Lasiosphaeridaceae based on Lasiosphaeris.
Lasiosphaeris Clem., Gen. fung. (Minneapolis): 35 (1909)
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, solitary, superficial to semi-immersed, subglobose to
obpyriform, black, ostiolate, tuberculate or/and surrounded
by brown, septate setae or hairs. Asci 8-spored, unitunicate,
cylindrical, with J- apical ring and a subapical globule.
Ascospores cylindrical to geniculate or sigmoid, aseptate to
multi-septate, ends rounded, hyaline to pale brown, becom-
ing brown to dark brown, ellipsoidal to broadly fusiform at
apical cell(s), sometimes with terminal, short, conical gelati-
nous appendage at each end. Asexual morph: Hyphomy-
cetous, phialophora-like. Conidia oval to globose, hyaline
(adapted from Tode 1791; Miller and Huhndorf 2004a).
Notes: Lasiosphaeris was established based on Sphaeria
hispida (Clements 1909) and Réblová etal. (2011) intro-
duced Lasiadelphia lasiosphaeriae for the asexual morph of
Lasiosphaeris hispida. However, Réblová etal. (2016) rec-
ommended to use the old Lasiosphaeris over Lasiadelphia.
Lasiosphaeris is characterized by hairy ascomata, cylindri-
cal to clavate asci with apical globule and geniculate or
Fungal Diversity
1 3
sigmoid ascospores (Tode 1791; Hilber and Hilber 1979),
which is related to Immersiella and Lasiosphaeria in mor-
phology, but distinct in phylogenetic analysis. Lasiospha-
eris species clustered as an independent clade in Sordariales
based on single/multi-gene analysis (Miller and Huhndorf
2004a; Kruys etal. 2014; Marin-Felix etal. 2020). This
resulted in Lasiosphaeris being placed in Sordariales genera
incertae sedis (Miller and Huhndorf 2004a; Wijayawardene
etal. 2020). Since a large number of lasiosphaeriaceous
taxa have been repositioned according to the phylogenetic
analysis (Marin-Felix etal. 2020), we raised Lasiosphaeris
as a novel family, Lasiosphaeridaceae, which includes all
Lasiosphaeris species and is branched out of other fami-
lies in Sordariales (72%ML) in our phylogenetic analysis
(Fig.26).
Type species: Lasiosphaeris hispida (Tode) Clem., Gen.
fung. (Minneapolis): [173] (1909)
Basionym: Sphaeria hispida Tode, Fung. mecklenb. sel.
(Lüneburg) 2: 17 (1791)
Facesoffungi number: FoF 10021; Fig.38
Saprobic on wood. Sexual morph: Ascomata
440 × 600µm (x̄ = 520µm, n = 10), perithecial, solitary or
gregarious, superficial to semi-immersed, globose to sub-
globose, black, membranaceous, ostiolate, with septate,
brown, tapering hairs, 3.5–5.5 wide. Peridium 45–75µm
(x̄ = 62 µm, n = 30) wide, membranaceous, compris-
ing two layers, outer layer composed of brown cells of
textura angularis; inner layer composed of hyaline cells
of textura prismatica. Paraphyses 2.5–5µm wide, numer-
ous, filiform. Asci (185–)210–230(–250) × 14–20 µm
(x̄ = 220 × 17µm, n = 30), 8-spored, unitunicate, cylindri-
cal, pedicellate, apex rounded, with apical globule, api-
cal ring distinct. Ascospores 60–70(–75) × 4.5–7.5µm
(x̄ = 65 × 6µm, n = 50), bi-seriate, cylindrical to genic-
ulate, slight curved near the base, hyaline and aseptate
when young, becoming pale brown and multi-septate when
mature, with a large guttule in each cell, smooth-walled.
Asexual morph: Undetermined.
Material examined: USA, Michigan, Marquette, Huron
Mountain Club, around Ives Lake, 45º 0.00ʹ N, 87º 0.00ʹ W,
on decayed wood, 17 August 1997, S.M. Huhndorf and M.H.
Huhndorf (F-SMH3336).
Known hosts and distribution: On dead wood in Germany
(type locality) and USA (Fuckel 1870; Miller and Huhndorf
2005).
Notes: The molecular data of Lasiosphaeris hispida were
provided (Miller and Huhndorf 2005; Vu etal. 2019). In this
study, L. hispida is basal to L. arenicola and L. hirsute in
Lasiosphaeridaceae (100%ML/1.00PP, Fig.26). We could
not obtain the type material. Therefore, we re-examined an
authentic specimen collected by Huhndorf.
New combination:
Lasiosphaeris arenicola (R. Hilber) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Cercophora arenicola R. Hilber, in Hilber &
Hilber, Z. Mykol. 45(2): 214 (1979)
Index Fungorum number: IF558252
Podosporaceae X. Wei Wang & Houbraken, in Wang etal.,
Stud. Mycol. 93: 223 (2019)
Coprophilous or saprobic on wood or isolated from soil.
Sexual morph: Ascomata perithecial or cleistothecial, soli-
tary to scattered or gregarious, immersed to semi-immersed
or superficial, obpyriform or ovoid, membranaceous to car-
bonaceous, brown to black, glabrous or surrounded by hairs.
Paraphyses cylindrical, septate. Asci 8-spored, unitunicate,
cylindrical to clavate, pedicellate, evanescent. Ascospores
uni- or bi-seriate, ovoid, ellipsoidal or triangular, 0–1-sep-
tate, hyaline to dark brown, with an apical germ pore.
Asexual morph: Hyphomycetous. Conidia globose to oval,
hyaline, smooth-walled, ovate masses on the apex of the
phialides (adapted from Wang etal. 2019a, b).
Type genus: Podospora (Corda) Ces. 1856
Notes: Three redefined genera, Cladorrhinum, Podospora
and Triangularia were classified in Podosporaceae based on
phylogenetic analyses (Wang etal. 2019a, b). Apiosordaria
was synonymized under Triangularia because its generic
type, A. verruculosa, was nested in Triangularia clade
(Wang etal. 2019a, b). In this study, species of Cladorrhi-
num, Podospora and Triangularia (= Apiosordaria) are scat-
tered in Sordariales, especially Podospora (Fig.26). These
three genera are polyphyletic with different characteristics;
Cladorrhinum has ornamented ascospores, Podospora has
ascospores with unique gelatinous appendages, while Trian-
gularia has polygonal ascospores (Krug etal. 1983; Guarro
and Cano 1988; Bell and Mahoney 1997). Herein we review
each genus of Podosporaceae based on phylogenetic analy-
ses and characteristics of the generic types.
Cladorrhinum Sacc. & Marchal, in Marchal, Bull. Soc. R.
Bot. Belg. 24(1): 64 (1885)
Saprobic on wood or isolated from soil, coprophilous.
Sexual morph: Ascomata perithecial, solitary to scattered,
immersed to superficial, globose to subglobose, semi-
transparent, brown to black, surrounded by hyaline, septate
hyphae. Peridium membranaceous, comprising pale brown
to reddish brown cells of textura intricata to textura epider-
moidea. Asci 8-spored, unitunicate, cylindrical to clavate to
obovoid, pedicellate, apex rounded, evanescent. Ascospores
ovoid to ellipsoidal, aseptate, hyaline to dark brown, with
an apical germ pore and a gelatinous appendage at one or
each end. Asexual morph: Hyphomycetous. Conidiophores
Fungal Diversity
1 3
Fig. 38 Lasiosphaeris hispida: (F-SMH 3336). a Herbarium mate-
rial label. b Herbarium material. c, d Ascomata on host. e Ascoma
in cross section. f Peridium. g–j Asci. k–n Ascospores. Notes: Fig
n soaked in Melzer's reagent. Scale bars: d = 500 µm, e = 200 µm,
f–j = 100µm, k–n = 20µm
Fungal Diversity
1 3
micronematous. Conidiogenous cells intercalary or terminal,
with a flared collarette, producing blastic conidia. Conidia
globose to oval, hyaline, aseptate, smooth-walled, with a
truncate base (adapted from Mouchacca and Gams 1993;
Wang etal. 2019a, b).
Notes: The hyphomycete Cladorrhinum was reported
as an asexual morph of Apiosordaria and Cercophora
(Mouchacca and Gams 1993), and it is characterized by clus-
tered dematiaceous hyphae, with a lateral phialide produc-
ing globose, aseptate, hyaline conidia. Cladorrhinum was
transferred to Podosporaceae based on multi-gene analysis
of the generic type (Wang etal. 2019a, b). Subsequently,
Marin-Felix etal. (2020) modified Cladorrhinum and trans-
ferred Arnium tomentosum and Cercophora coprophila as
Cla. tomentosum and Cla. coprophilum respectively based
on phylogenetic analysis. In this study, four species are
transferred to Cladorrhinum based on their molecular data
(see the combination list below, Fig.26).
Type species: Cladorrhinum foecundissimum Sacc. & Mar-
chal, in Marchal, Bull. Soc. R. Bot. Belg. 24(1): 64 (1885)
Facesoffungi number: FoF 10127; Fig.37d
Coprophilous or isolated from soil. Sexual morph:
Undetermined. Asexual morph: Hyphomycetous. Mycelium
yellowish-grey, velvety, dense. Hyphae filiform, septate,
hyaline, branched. Conidiophores micronematous. Conid-
iogenous cells intercalary, with a flared collarette, producing
blastic conidia. Conidia globose, hyaline, smooth-walled,
with a truncate base, masses on the apex of the phialides
(adapted from Marchal 1885; Wang etal. 2019a, b).
Known hosts and distribution: On wild boar dung in Bel-
gium (type locality) (Marchal 1885); on soil in the Nether-
lands (Wang etal. 2019a, b).
Notes: Hyphomycetous Cladorrhinum foecundissimum
was first isolated from wild boar dung in Belgium (Marchal
1885). Wang etal. (2019a, b) provided an ex-neotype strain
for Cla. foecundissimum (CBS 180.66) which was isolated
from soil in the Netherlands since the holotype was lost. In
this study, we found that this strain is sister to Cla. gran-
diusculum and Cla. olerum with 96%ML/0.93PP support
(Fig.26).
New combinations:
Cladorrhinum grandiusculum (A.E. Bell & Mahoney)
S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Cercophora grandiuscula A.E. Bell &
Mahoney, in Bell, CBS Diversity Ser. (Utrecht) 3: 59 (2005)
Index Fungorum number: IF558255
Cladorrhinum leucotrichum (Speg.) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Sordaria leucotricha Speg., Michelia 1(no.
2): 225 (1878)
Synonym: Zopfiella leucotricha (Speg.) Malloch & Cain,
Can. J. Bot. 49(6): 876 (1971)
Index Fungorum number: IF558256
Cladorrhinum terricola (S. Ueda) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Cercophora terricola S. Ueda, Mycoscience
35(3): 287 (1994)
Index Fungorum number: IF558257
Cladorrhinum olerum (Fr.) S.K. Huang & K.D. Hyde,
comb. nov.
Basionym: Sphaeria olerum Fr., Elench. fung. (Greif-
swald) 2: 98 (1828)
Synonym: Arnium olerum (Fr.) N. Lundq. & J.C. Krug,
Symb. bot. upsal. 20(no. 1): 212 (1972)
Facesoffungi number: FoF 10022; Index Fungorum num-
ber: IF558330; Fig.39
Coprophilous, saprobic on herbaceous plant. Sexual
morph: Ascomata perithecial, scattered, semi-immersed,
broadly obpyriform to subglobose, membranaceous, pale
brown to dark brown, semi-transparent, ostiolate, with a
black papilla, periphysate. Peridium composed of yellowish
cells of textura intricata. Asci 8-spored, unitunicate, cylin-
drical, pedicellate, apex rounded, evanescent. Ascospores
(40–)47–58(–65) × 20–40µm (x̄ = 50 × 30µm, n = 20), uni-
seriate, obovoid to ellipsoidal, aseptate, hyaline when young,
becoming dark brown when mature, with ornamented wall
and a gelatinous appendage at each end. Asexual morph:
Undetermined.
Material examined: Norway, Nord-Trøndelag,
Grong, Godejorden, on old cow dung after 1–2months
in moist chamber in Uppsala, 17 June 1962, N. Lun-
dqvist (UPS-UPS:BOT:F-125924); Sweden, Gotland,
Visby par., Skansudd, NE of Visby, on old cow dung in
moist chamber in Uppsala, 30 July 1961, N. Lundqvist
(UPS- UPS:BOT:F-125925).
Known hosts and distribution: On stems of Brassica oler-
acea in Franch (type locality) (Fries 1828); usually on cab-
bage stalk in Austria, Belgium, Czech Republic, Denmark,
England, Finland, France, Germany, Morocco, Norway,
Portugal, Sweden (Lundqvist 1972; Jeng and Krug 1977;
Kruys etal. 2014).
Notes: We re-examined two specimens of Cladorrhinum
olerum which was collected and determined by Lundqvist
(1972), and the hand-drawing of ascus and ascospores are
provided (Fig.39e–f) based on Kruys etal. (2014). Phy-
logenetically, Arnium olerum nested in Lasiosphaeriaceae
(Kruys etal. 2014) and was transferred to Cladorrhinum
(Marin-Felix etal. 2020). In this study, Cla. olerum is sister
to Cla. grandiusculum in Podosporaceae (58%ML, Fig.26).
Fungal Diversity
1 3
Podospora Ces., Hedwigia 1(15): 103 (1856)
Synonym: Schizothecium Corda, Icon. fung. (Prague) 2:
29 (1838)
Saprobic on wood or isolated from soil. Sexual morph:
Ascomata perithecial, solitary to scattered, immersed to
semi-immersed, obpyriform to ovoid, brown to black, ostio-
late. Peridium membranaceous to coriaceous, composed of
pale brown to reddish brown-walled cells of textura por-
recta to textura angularis to textura prismatica. Paraphyses
cylindrical, septate. Asci 8-spored, unitunicate, cylindrical to
clavate, pedicellate, apex rounded, evanescent. Ascospores
ovoid to ellipsoid, aseptate, hyaline when young, becoming
a swollen, dark brown upper cell and a hyaline lower cell
when mature, with a gelatinous appendage at one or each
end. Asexual morph: Hyphomycetous, cladorrhinum-
like. Conidiophores micronematous. Conidiogenous cells
intercalary or terminal, with a flaring collarette producing
blastic conidia. Conidia globose to oval, aseptate, hyaline,
smooth-walled, with a truncate base (adapted from Cesati
1856; Lundqvist 1972).
Notes: Podospora is typified by P. fimicola which has
coriaceous ascomata covered with simple short hairs and
ascospores with gelatinous appendages (Cesati 1856). Lun-
dqvist (1972) clarified Podospora species and proposed
Fig. 39 Cladorrhinum olerum–as Arnium olerum: a (UPS-
UPS:BOT:F-125924); b–d, g–l (UPS-UPS:BOT:F-125925); e,
f (redrawn from Kruys et al. 2014). a, b Squashed ascomata c
Peridium. d, e Asci. f–l Ascospores. Notes: a–d, g–l from perma-
nent mount. Scale bars: a–b = 500 µm, c. e = 50 µm, d = 100 µm,
f–l = 20µm
Fungal Diversity
1 3
that Podospora fimiseda is a name change for Podospora
fimicola. Thus, these species are conspecific (Niessl 1883;
Lundqvist 1972; Bell and Mahoney 1997; Doveri 2008).
However, following priority, Wang etal. (2019a, b) accepted
P. fimicola as its generic type and redefined Podospora, the
type genus of Podosporaceae. Phylogenetically, we transfer
seven species clustered in Podospora clade, to Podospora
to avoid confusion (see the combination list below, Fig.26).
Species of Podospora (s. lat.) are also scattered in Bombar-
diaceae, Lasiosphaeriaceae, Naviculisporaceae and Neoschi-
zotheciaceae (Fig.26).
Type species: Podospora fimicola (Corda) Ces., Hedwigia
1(15): 103 (1856)
Basionym: Schizothecium fimicola Corda [as 'fimicolum'],
Icon. fung. (Prague) 2: 29 (1838)
Facesoffungi number: FoF 10128; Fig.33a–d
Coprophilous. Sexual morph: Ascomata perithecial,
solitary to scattered, immersed to semi-immersed, obpyri-
form to ovoid, membranaceous, brown to black, ostiolate.
Peridium comprising three layers, outer layer composed of
pale brown to reddish brown-walled cells of textura por-
recta to textura angularis; inner layer composed of hyaline
cells of textura prismatica. Paraphyses cylindrical, septate.
Asci 8-spored, unitunicate, cylindrical to clavate, pedicel-
late, apex rounded, evanescent. Ascospores bi-seriate, ovoid
to ellipsoid, aseptate, hyaline when young, becoming dark
brown when mature, with ornamented wall and a gelatinous
appendage at each end, with longitudinally striate. Asexual
morph: Hyphomycetous (Cladorrhinum on CMA). Conidia
globose to oval, hyaline, smooth-walled (adapted from Bell
and Mahoney 1997).
Material examined: Italy, Piemonte, Biella, Bugellensi-
bus Mountain, on cow dung, 1855, V. Cesati (S-F26041,
lectotype).
Known hosts and distribution: On dry cattle dung on dry
pastures in Czech Republic (type locality) (Corda 1838); on
cow, horse dung in Austria, Belgium, Brazil, Canada, Chile,
China, Denmark, Finland, France, Germany, Hungary, Italy,
Liberia, Norway, Poland, Puerto Rico, Romania, Russia,
Scotland, Spain, Sweden, USA, and Venezuela (Lundqvist
1972).
Notes: Lundqvist (1972) reviewed samples of related
Podospora fimicola and found that this species is generally
found in the faeces of herbivores. Vu etal. (2019) sequenced
the epitype strain of Schizothecium fimicola (= Podospora
fimicola), CBS 482.64. This species is sister to P. costari-
censis in Podosporaceae (80%ML, Fig.26). In this study,
we observed the lectotype material of P. fimicola however
only the dried ascomata could be seen. We drew the ascus
and ascospores of P. fimicola concerning Bell and Mahoney
(1997) and Doveri (2008).
New combinations:
Podospora brunnescens (W. Gams) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Cladorrhinum brunnescens W. Gams, Myco-
taxon 48: 435 (1993)
Index Fungorum number: IF558259
Podospora dacryoidea S.K. Huang & K.D. Hyde, nom. nov.
Synonym: Cladorrhinum australe Gasoni, in Carmarán,
Berretta, Martínez, Barrera, Munaut & Gasoni, Mycol.
Progr. 14(no. 94): 9 (2015)
Index Fungorum number: IF558260
Etymology: ‘dacryoidea’ refers to Carmarán etal. (2015)
mentioned that this fungus has dacryoid conidia.
Podospora flexuosa (Madrid etal.) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Cladorrhinum flexuosum Madrid, Cano, Gené
& Guarro, Mycologia 103(4): 801 (2011)
Index Fungorum number: IF558261
Podospora jamaicensis (B.M. Robison) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Echinopodospora jamaicensis B.M. Robison,
Trans. Br. mycol. Soc. 54(2): 318 (1970)
Synonym: Apiosordaria jamaicensis (B.M. Robison) J.C.
Krug, Udagawa & Jeng, Mycotaxon 17: 546 (1983)
Index Fungorum number: IF558298
Podospora hamata (Wu etal.) S.K. Huang & K.D. Hyde,
comb. nov.
Basionym: Apiosordaria hamata B. Wu, K.D. Hyde, Jing
Z. Sun & Xing Z. Liu, in Wu, Tian, Wang, Liu, Hyde & Sun,
Mycotaxon 131(4): 852 (2016) [2017]
Index Fungorum number: IF558299
Podospora macrospora (Guarro & Calvo) S.K. Huang &
K.D. Hyde, comb. nov.
Basionym: Zopfiella macrospora Guarro & Calvo, Nova
Hedwigia 37(4): 663 (1983)
Index Fungorum number: IF558300
Podospora spinosa (Cailleux) S.K. Huang & K.D. Hyde,
comb. nov.
Basionym: Tripterospora spinosa Cailleux, Cahiers de La
Maboké 8(1): 16 (1970)
Synonym: Apiosordaria spinosa (Cailleux) J.C. Krug,
Udagawa & Jeng, Mycotaxon 17: 546 (1983)
Index Fungorum number: IF558301
Triangularia Boedijn, Annls mycol. 32(3/4): 302 (1934)
Synonym: Apiosordaria Arx & W. Gams, Nova Hedwigia
13: 201 (1967)
Fungal Diversity
1 3
Isolated from soil. Sexual morph: Ascomata cleistothe-
cial or perithecial, gregarious to scattered, superficial to
immersed, membranaceous or carbonaceous or coriaceous,
obpyriform to pyriform, black, opaque or semi-transparent,
glabrous or surrounded by hairs or setae, ostiolate, with
papilla. Peridium comprising membranaceous to coria-
ceous, pale brown to reddish brown cells. Paraphyses fili-
form, septate. Asci 4–8–multi-spored, unitunicate, clavate
to cylindrical, pedicellate, evanescent. Ascospores ellipsoi-
dal to triangular, aseptate, hyaline when young, becoming
1-septate, upper cell larger, brown, ellipsoidal to polygo-
nal, smooth-walled or with ornamented wall, with numer-
ous broad or narrow pits, with a sub-apical or apical germ
pore; lower cell smaller, hyaline, sometimes collapsing, with
gelatinous appendages at one or each end, smooth-walled or
sometimes with ornamented on surface, guttulate. Asexual
morph: Hyphomycetous, cladorrhinum-like. Conidiophores
micronematous. Conidiogenous cells intercalary or terminal,
with a flaring collarette producing blastic conidia. Conidia
globose to oval, aseptate, hyaline, smooth-walled, with a
truncate base (adapted from Boedijn 1934; Guarro and Cano
1988; Wang etal. 2019a, b).
Notes: Triangularia was transferred from Trigonia and, is
typified by T. bambusae in having glabrous ascomata or those
covered with hairs and ellipsoidal to triangular ascospores
(Boedijn 1934; Guarro and Cano 1988). This genus was pre-
viously accepted in Lasiosphaeriaceae (Wijayawardene etal.
2018), and Wang etal. (2019a, b) redefined Triangularia as
a member of Podosporaceae based on multi-gene analysis.
Apiosordaria was synonymized under Triangularia, because
the strain of its generic type A. verruculosa (CBS 148.77)
(Fig.33i–j) nested in Triangularia (Wang etal. 2019a, b).
Forin etal. (2021) accepted Triangularia in Lasiospha-
eriaceae. In this study, seven species clustered in Triangu-
laria were transferred to Triangularia to avoid confusion (see
the combination list below, Fig.26). Species of Apiosordaria
and Triangularia are also scattered in Naviculisporaceae and
Neoschizotheciaceae in the phylogenetic analysis (Fig.26).
Type species: Triangularia bambusae (J.F.H. Beyma)
Boedijn, Annls mycol. 32(3/4): 302 (1934)
Basionym: Trigonia bambusae J.F.H. Beyma, Centbl.
Bakt. ParasitKde, Abt. II 89: 236 (1933)
Facesoffungi number: FoF 10129; Fig.37e.
Isolated from soil. Sexual morph: Ascomata perithecial,
gregarious to scattered, superficial to immersed, ovoid to
pyriform, black, glabrous or surrounded by hairs. Peridium
comprising membranaceous, pale brown to reddish brown
cells of textura angularis. Asci 8-spored, unitunicate, clavate
to cylindrical, pedicellate, evanescent. Ascospores uni- or bi-
seriate, ellipsoidal to triangular, aseptate and hyaline when
young, becoming conical with an oblique septum near one
end, upper cell triangular, brown, with a subapical germ
pore; lower cell irregular or hemispherical, hyaline, collaps-
ing. Asexual morph: Undetermined (adapted from Guarro
and Cano 1988).
Known hosts and distribution: On culms of Bambusaceae
in Netherlands (type locality) (Van Beyma Thoe Kingma
1933).
Notes: Triangularia bambusae is sister to T. batistae in
Podosporaceae based on phylogenetic analysis (Vu etal.
2019; Wang etal. 2019a, b; this study, 100%ML/1.00PP,
Fig.26).
New combinations:
Triangularia microsclerotigena (Madrid etal.) S.K.
Huang & K.D. Hyde, comb. nov.
Basionym: Cladorrhinum microsclerotigenum Madrid,
Cano, Gené & Guarro, Mycologia 103(4): 804 (2011)
Index Fungorum number: IF558302
Triangularia nannopodalis (Cain) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Podospora nannopodalis Cain, Can. J. Bot.
40(3): 455 (1962)
Index Fungorum number: IF558303
Triangularia praecox (Cailleux) S.K. Huang & K.D. Hyde,
comb. nov.
Basionym: Podospora praecox Cailleux, Cahiers de La
Maboké 7: 102 (1969)
Index Fungorum number: IF558304
Triangularia samala (Udagawa & T. Muroi) S.K. Huang &
K.D. Hyde, comb. nov.
Basionym: Cercophora samala Udagawa & T. Muroi,
Trans. Mycol. Soc. Japan 20(4): 454 (1979)
Index Fungorum number: IF558305
Triangularia tarvisina (Sacc.) S.K. Huang & K.D. Hyde,
comb. nov.
Basionym: Philocopra setosa subsp. tarvisina Sacc., Syll.
fung. (Abellini) 1: 250 (1882)
Synonym: Podospora tarvisina (Sacc.) J.H. Mirza &
Cain, Can. J. Bot. 47(12): 2041 (1970) [1969]
Index Fungorum number: IF558306
Triangularia unicaudata (C. Moreau & M. Moreau ex G.
Sm.) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Pleurage unicaudata C. Moreau & M. Moreau
ex G. Sm., Trans. Br. mycol. Soc. 40(4): 488 (1957)
Synonym: Podospora unicaudata (C. Moreau & M.
Moreau ex G. Sm.) Cain, Can. J. Bot. 40: 460 (1962)
Index Fungorum number: IF558321
Fungal Diversity
1 3
Triangularia yaeyamensis (Morinaga etal.) S.K. Huang &
K.D. Hyde, comb. nov.
Basionym: Apiosordaria yaeyamensis Morinaga, Utatsu
& Minoura, Trans. Mycol. Soc. Japan 20(4): 493 (1979)
Index Fungorum number: IF558322
Sordariaceae G. Winter [as 'Sordarieae'], Rabenh. Krypt.-
Fl., Edn 2 (Leipzig) 1.2: 162 (1885)
Saprobic or endophytic on herbs or woody plants, isolated
from soil or/and dung, in terrestrial and/or aquatic habitats.
Sexual morph: Ascomata perithecial or cleistothecial,
solitary, scattered or gregarious, superficial, erumpent or
immersed, globose, subglobose, pyriform to ovoid, car-
bonaceous, coriaceous or membranaceous, yellow, oliva-
ceous brown, brown to black, tuberculate or glabrous, with
setae or not, ostiolate or not, with papilla or indistinct or
papilla absent, with hyaline periphyses or periphyses absent.
Peridium comprising hyaline to brown-walled cells, carbo-
naceous, coriaceous or membranaceous. Paraphyses numer-
ous, branched or unbranched, septate, filiform or cylindrical,
evanescent, or absent. Asci 2–8-spored, unitunicate, cylindri-
cal to clavate, pedicellate, with J- apical ring or ring indis-
tinct or absent. Ascospores uni–bi-seriate, hyaline, becom-
ing yellowish or olivaceous brown or brown or black when
mature, globose to ellipsoidal, aseptate, with germ pore(s)
or absent, smooth-walled or with angular to circular reticu-
late patterns or pits or longitudinal ribs on surface, with a
sheath or sheath absent, guttulate. Asexual morph: Hypho-
mycetous. Hyphae flexuous, hyaline to dark brown, septate,
branched. Conidiophores hyaline to brown, branched or
unbranched, straight or flexuous, smooth-walled, septate.
Conidia hyaline, yellow, orange or dark brown, oval to
ellipsoidal, smooth-walled, with a sheath or sheath absent
(adapted from Lundqvist 1972; García etal. 2004; Cai etal.
2006b; Maharachchikumbura etal. 2016).
Type genus: Sordaria Ces. & De Not. 1863
Notes: Sordariaceae was established by Winter (1885), and
traditionally eight genera, Copromyces, Effetia, Gelasinos-
pora, Guilliermondia, Neurospora, Pseudoneurospora, Sor-
daria and Stellatospora have been accommodated (Maharach-
chikumbura etal. 2015, 2016; Wijayawardene etal. 2018).
Most of the species in this family grow on dung, soil or decay-
ing substrates in terrestrial and/or aquatic habitats (Frederick
etal. 1969; Lundqvist 1972; Guarro and von Arx 1987; Krug
and Scott 1994; García etal. 2004; Cai etal. 2006b), and they
are closely related to Lasiosphaeriaceae, based on phyloge-
netic analysis and similarities in perithecial ascomata (Lun-
dqvist 1972; Huhndorf etal. 2004b; Maharachchikumbura
etal. 2015, 2016; Fig.1, 26). Due to the typical morphology
of ascospores, the genera in Sordariaceae have undergone
various taxonomic changes. The monotypic genera, Effetia
and Guilliermondia were included in Sordariaceae because
of their characteristics are similar to Sordaria (Boudier 1904;
Lundqvist 1967; Bartoli etal. 1984; Maharachchikumbura
etal. 2015). Copromyces lacked reliable molecular data and
was suspected of resembling Rechingeriella (Dothideomy-
cetes) (Petrak 1967). Thus, we recommend placing this genus
in Ascomycota genera incertae sedis pending future work.
Pseudoneurospora was independent of Neurospora because
it has unique ascospores with irregular ornamentation on
the surface and it also exists as a separate cluster in molecu-
lar analysis (García etal. 2004). The monotypic Boothiella
was accepted in Sordariaceae based on its type strain (CBS
334.67) nested in Sordariaceae (Wang etal. 2019a, b; this
study). Wang etal. (2019a, b) removed Stellatospora from
Sordariaceae as its ex-type strain, Stellatospora terricola
(CBS 811.95), clusters in Chaetomiaceae. Gelasinospora
was synonymized under Neurospora based on their similar
morphology and close phylogenetic relationship (García etal.
2004). The traditional genera Neurospora and Sordaria were
previously defined by morphology: Neurospora, which means
‘nerve spore’ refers to the ascospores with axon-like ornamen-
tation on the surface (Shear and Dodge 1927; García etal.
2004). Sordaria is a genus mainly found in the faeces and has
ascospores with sheaths (Cesati and de Notaris 1863; Maha-
rachchikumbura etal. 2016). These two genera are polyphy-
letic in Sordariales (Spatafora and Blackwell 1994; Dettman
etal. 2001; García etal. 2004; Cai etal. 2006b; this study). In
this study, we found that all Neurospora and Sordaria species
clustered in Sordariaceae. Their respective positions are usu-
ally unstable in multi-gene (Fig.40) or single-gene analyses
(not shown). Hence, Neurospora and Sordaria can be mainly
distinguished by morphology. Neurospora is characterized by
globose to pyriform, brown to black ascomata, cylindrical to
broadly clavate asci and aseptate ascospores with a variety of
ribs or pits, whereas, Sordaria has subglobose to ovoid, brown
to black ascomata, cylindrical to clavate asci and ellipsoidal to
ovoid ascospores with smooth-walls and may or may not have
sheath. However, in this study, we include five genera in Sor-
dariaceae based on morphology and phylogenetic analyses.
Boothiella Lodhi & Mirza, Mycologia 54(2): 217 (1962)
Isolated from soil. Sexual morph: Ascomata cleistothe-
cial, solitary or gregarious, superficial to immersed in the
medium, globose to subglobose, membranaceous, brown to
black, glabrous. Peridium composed of membranaceous, hya-
line to subhayline cells of textura angularis. Asci 4-spored,
unitunicate, clavate to cylindrical, short pedicellate, apex
rounded, evanescent. Ascospores hyaline, becoming dark
brown, oval to citriform, 1–2-apiculate, aseptate, with a germ
pore at one or each end. Asexual morph: Undetermined
(adapted from Lodhi and Mirza 1962; Wang etal. 2019a, b).
Notes: The monotypic genus Boothiella was introduced
as a member of Eurotiales and its ascospores are similar to
Thielavia (Lodhi and Mirza 1962). Kirk etal. (2008) trans-
ferred Boothiella to Sordariaceae and Maharachchikumbura
Fungal Diversity
1 3
0.04
Sordaria alcina CBS 109462
Sordaria lappae CBS 154.97
Neurospora tetrasperma CBS 223.38
Sordaria islandica CBS 512.77 T
Phialemonium obovatum CBS 279.76 T
Neurospora intermedia CBS 131.92
Neurospora africana CBS 571.69 T
Sordaria conoidea CBS 563.72
Phialemonium atrogriseum CBS 604.67 T
Sordaria fimicola CBS 511.77
Neurospora galapagosensis FGSC 1739
Neurospora udagawae CBS:309.91 T
Phialemonium limoniforme CBS 139049 T
Neurospora hippopotama CBS:561.94 T
Sordaria alcina CBS 109460
Neurospora saitoi CBS:435.74 T
‘Copromyces octosporus’ CBS 386.78 T
Neurospora novoguineensis CBS 495.81 T
Sordaria macrospora CBS 346.62
Neurospora tetraspora CBS:178.33 T
Neurospora reticulata CBS 435.61 T
Neurospora retispora CBS 275.50 T
Sordaria sibutii CBS 568.72 T
Neurospora perkinsii FGSC 8838
Neurospora foveaconica CBS 557.72 T
Chaetomium globosum CBS 160.62 T
Neurospora calospora CBS 444.78
Neurospora inversa CBS 554.72 T
Neurospora longispora CBS 142.68 T
Sordaria mabokeensis CBS 566.72 T
Chaetomium tenue CBS 139.38 T
Neurospora kobi CBS 560.72 T
Lasiosphaeria glabrata SMH4617
Sordaria brevicollis FGSC 1904 T
Pseudoneurospora amorphoporcata CBS 626.80
Neurospora cerealis CBS 256.52
Neurospora terricola CBS 298.63 T
Neurospora pseudoreticulata CBS 556.72 T
Neurospora lineolata CBS:502.70 T
Boothiella tetraspora CBS 334.67 T
Sordaria goundaensis var. latispora CBS 565.72 T
Sordaria tamaensis NBRC 32552 T
Neurospora sitophila CBS 112.19
Sordaria fimicola CBS 508.50
Lasiosphaeria lanuginosa SMH3819
Neurospora crassa ICMP 6360
Neurospora brevispora CBS:547.94 T
Neurospora dictyophora CBS:529.95 T
Sordaria tomentoalba CBS 569.72 T
Chaetomium pseudoglobosum CBS 574.71 T
Neurospora santi-florii CBS:571.72 T
Neurospora bonaerensis CBS 102191
Gelasinospora autosteira CBS 265.51 T
Neurospora indica FGSC 7793
Lasiosphaeria sorbina CBS 885.85
Sordaria prolifica CBS 567.72 T
Sordaria arctica CBS 143.68
Neurospora calospora IFO 32008
Neurospora sublineolata IMI 223881 T
Neurospora pannonica TRTC 51327 T
Boothiella tetraspora CBS 887.97
Pseudoneurospora canariensis CBS 135818 T
Sordaria nodulifera NBRC 32551 T
Neurospora endodonta IMI 148369 T
Neurospora hispaniola FGSC 8817 T
Neurospora hapsidophora CBS 197.95 T
Neurospora stellata CBS 561.72 T
Lasiosphaeria ovina CBS 126299
Neurospora heterospora CBS 559.72 T
Sordaria superba CBS 784.96
Sordaria sclerogenia FGSC 2741
Neurospora nigeriensis FMR 5963 T
Neurospora seminuda CBS:572.72 T
Neurospora uniporata FMR 7283 T
Neurospora cratophora CBS:558.94 T
Sordaria goundaensis CBS 564.72 T
Neurospora metzenbergii FGSC 8847
Neurospora varians CBS 668.77 T
Neurospora micropertusa IMI 212194
Neurospora dodgei FGSC 1692
Neurospora discreta FGSC 3228 T
Neurospora goundaensis CBS 558.72 T
Sordaria humana CBS 248.89
53/0.95
100/1.00
55/--
99/1.00
54/--
51/0.93
93/0.98
94/0.99
93/0.99
59/--
59/--
62/0.97
92/1.00
56/0.93
80/--
100/1.00
83/1.00
62/--
100/1.00
100/1.00
85/0.99
94/0.90
91/--
100/1.00
87/1.00
60/--
97/1.00
100/1.00
66/--
87/0.93
100/1.00
67/0.98
80/1.00
51/--
75/0.95
63/0.93
100/1.00
100/1.00
78/1.00
Mating type:
Heterothallic
Pseudohomothallic
Homothallic
HE
PS
HO
Ascospores appearance:
Pitted
Ribbed
Reticulated
Smooth-walled
with sheath
PI
RI
RE
SM
SH
Form of Ascomata:
Perithecium
Cleistothecium
PE
CL
Types of epispore:
Ornate
Smooth
Epispore
HO PE PI
Mating type Ascomata Ascospores
HO PE/CL PI
HO PE RE
HO PE RE
HO CL PI
PE/CLPI
HO PE PI
HO PE PI
HO CL PI
PS PE PI
HO CL RI&PI
HO PE RI
HE PE SM&SH
HO PE SM
HO PE SM
HO PE SM
HO PE SM
HE PE SM&SH
HE PE SM&SH
HO PE SM
HO PE SM
HE PE RI
HE
HE
HE
HE PE RI
HE PE
RI
PS PE
RI&PI
HE PE RI&PI
HO PE SM
HO PE SM&SH
PE SM
PE SM&SH
HO PE SM&SH
PE SM&SH
PE SM&SH
PE SM
HO PE SM&SH
HO PE SM&SH
HO PE RE
CL RE
PE SM&SH
PE SM&SH
HO PE/CL RE
HE PE PI
HO PE RI
CL PI
PE PI
PE PI
HO PE PI
PE PI
HO PE RE
HO PE PI&RE
HO PE PI&RE &
PE PI
HO CL RE
HO PE RE
HO PE RI
HO PE RI
HO PE PI
HO PE RI
HO PE RI
HO CL PI
HO PE PI
PE PI
HO CL RE
PE RE
HO PE PI
HO PE PI
HO CL RE
CL SM
CL SM
Neurospora I
Sordaria I
Neurospora II
Sordaria II
Pseudoneurospora
Sordaria III
Neurospora III
Neurospora IV
Boothiella
Sordariaceae
Chaetomiaceae
Lasiosphaeriaceae
Cephalothecales
Sordariales
--/0.93
--/0.94
&
&&
&
&
N. autosteira
?
Fig. 40 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS and TUB sequence data of Sordari-
aceae. The confidence values of bootstrap (BS) proportions from the
Maximum Likelihood (ML) analysis (ML-BS > 50%, before the back-
slash) and the posterior probabilities (PP) from the Bayesian (BY)
analysis (BY-PP > 0.90, after the backslash) above corresponding
nodes. The ‘--’ indicates lack of statistical support (< 50% for ML-BS
and < 0.90 for BY-PP). Related sequences were referred to Cai etal.
(2006b), Wang etal. (2019a, b) and Vu et al. (2019). Eighty-four
strains are included in the combined analyses which comprise 1909
characters (862 characters for LSU, 583 characters for ITS, 464
characters for TUB) after alignment. Strains of Cephalothecales are
used as the outgroup taxa. The model of each partitioned gene is
GTR + I + G. The best score in IQ-TREE explores with a final like-
lihood value of -9569.3117 is presented. Sequences generated indi-
cated in bold. Strain numbers are noted after the species names and
ex-type strains are marked with ‘T’ after the culture number. Some
important characters of Sordariaceae species treated in this study are
illustrated at the right side of the tree. Alignments are available at
TreeBASE (URL: http:// purl. org/ phylo/ treeb ase/ phylo ws/ study/ TB2:
S28269)
Fungal Diversity
1 3
etal. (2015, 2016) placed this genus in Chaetomiaceae. Vu
etal. (2019) and Wang etal. (2019a, b) analyzed sequence
data from isolates CBS 334.67 (ex-type) and CBS 887.97,
and showed that B. tetraspora nests in Sordariaceae. In the
present study, Boothiella strains form a clade in Sordari-
aceae (100%ML/1.00PP, Figs.26, 40).
Type species: Boothiella tetraspora Lodhi & J.H. Mirza,
Mycologia 54(2): 217 (1962)
Facesoffungi number: FoF 10130; Fig.68r–s
Isolated from soil. Sexual morph: Ascomata cleistothe-
cial, solitary or gregarious, superficial to immersed in the
medium, globose to subglobose, membranaceous, brown to
black, glabrous. Peridium composed of membranaceous, hya-
line to subhayline cells of textura angularis. Asci 4-spored,
unitunicate, clavate to cylindrical, short pedicellate, apex
rounded, evanescent. Ascospores hyaline, becoming dark
brown, oval to citriform, 1–2-apiculate, aseptate, with a germ
pore at one or each end. Asexual morph: Undetermined
(adapted from Lodhi and Mirza 1962; Wang etal. 2019a, b).
Known hosts and distribution: On soil in Pakistan (type
locality) (Lodhi and Mirza 1962); on sand in Spain (Wang
etal. 2019a, b).
Notes: Boothiella tetraspora is similar to Thielavia,
except for its translucent ascomata (Lodhi and Mirza 1962).
Wang etal. (2019a, b) proposed that this species resembles
Hyalosphaerella fragilis and Pseudothielavia subhyalo-
derma (Chaetomiaceae), but the former has 4-spored and
the latter two have 8-spored asci. However, B. tetraspora
nests in Sordariaceae based on phylogenetic analyses (Wang
etal. 2019a, b; this study, Fig.26, 40).
Guilliermondia Boud., Bull. Soc. mycol. Fr. 20: 19 (1904)
Coprophilous on horse dung. Sexual morph: Ascomata
cleistothecial, solitary or gregarious, superficial, dark brown
to black, tuberculate. Peridium outer layer composed of
carbonaceous, dark tissues; inner layer composed of mem-
branaceous, hyaline to dark brown cells of textura angula-
ris. Paraphyses numerous, evanescent, filamentous, septate,
branched. Asci 8-spored, unitunicate, spherical, without an
apical ring, with long and evanescent pedicel. Ascospores
overlapping, ellipsoidal, brown, aseptate, smooth-walled,
surrounded by hyaline or brown gelatinous sheath. Asexual
morph: Undetermined (adapted from Boudier 1904).
Notes: The monotypic Guilliermondia is typified by G.
saccoboloides has black ascomata and ellipsoidal ascospores
surrounded by a brown sheath (Boudier 1904). Initially,
Boudier (1904) proposed that this genus is related to Phil-
lipsiella (Saccardiaceae), and Guilliermondia was later con-
sidered similar to Sordaria and transferred to Sordariaceae
(Kirk etal. 2008; Lumbsch and Huhndorf 2010; Hyde etal.
2020). In this study, Guilliermondia is placed in Sordari-
aceae, because the generic type G. saccoboloides is similar
to Sordaria fimicola. Both taxa are isolated from the faeces
of herbivores and have dark ascomata and ellipsoidal, brown
ascospores with gelatinous sheaths.
Type species: Guilliermondia saccoboloides Boud., Bull.
Soc. mycol. Fr. 20: 20 (1904)
Facesoffungi number: FoF 10023; Fig.41
Coprophilous on horse dung. Sexual morph: Ascomata
295–460 × 170–275µm (x̄ = 350 × 230µm, n = 10), cleistothe-
cial, solitary or gregarious, superficial, dark brown to black,
tuberculate. Peridium outer layer composed of carbonaceous,
dark tissues; inner layer composed of membranaceous, hyaline
to dark brown cells of textura angularis. Paraphyses 1–2µm
wide, numerous, evanescent, filamentous, septate, branched.
Asci 20–30µm (x̄ = 25µm, n = 30) diam. at ascospore-bearing
part, 8-spored, unitunicate, spherical, without an apical ring,
with long and evanescent pedicel. Ascospores 10–16 × 6–9µm
(x̄ = 14 × 7µm, n = 50), overlapping, ellipsoidal, brown, asep-
tate, smooth-walled, surrounded by a hyaline or brown gelati-
nous sheath 1–2.5µm thick. Asexual morph: Undetermined
(adapted from Boudier 1904).
Material examined: France, Lyon, 69 Rhône, on horse
dung, April 1903, M.A.A. Guilliermond (PC-MNHN-PC-
PC0167691, holotype).
Known hosts and distribution: On horse dung in France
(type locality) (Boudier 1904).
Notes: We re-examined the holotype, MNHN-PC-
PC0167691, and found that most of the ascomata had
collapsed and the pedicel of the asci were basically
inconspicuous.
Neurospora Shear & B.O. Dodge, J. Agric. Res., Washing-
ton 34: 1025 (1927)
Coprophilous or isolated from soil, burned or unburned
vegetation in terrestrial habitats, heterothallic or pseudoho-
mothallic. Sexual morph: Ascomata perithecial or cleis-
tothecial, gregarious or scattered, solitary, semi-immersed
to superficial, globose, subglobose to ovoid, brown to
black, ostiole with hyaline periphyses. Peridium compris-
ing brown to hyaline cells of textura angularis, textura
globulosa, textura prismatica, textura intrincata or textura
epidermoidea, carbonaceous to membranaceous. Paraphy-
ses numerous, filiform or cylindrical or lacking. Asci 4- or
8-spored, unitunicate, cylindrical to clavate, pedicellate,
with J- apical ring. Ascospores hyaline, olivaceous brown,
grey or black, oval to ellipsoidal, aseptate, with longitudinal
ribs or irregular to circular, deep or shallow pits on surface
and germ pore(s) at one/each end, with or without guttules.
Asexual morph: Hyphomycetous. Mycelium hyaline to light
brown, septate. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells phialidic, oval to clavate, hyaline,
smooth-walled. Conidia ellipsoidal to oval, hyaline, yellow,
orange or dark brown, smooth-walled (adapted from Shear
Fungal Diversity
1 3
and Dodge 1927; Tai 1935; Perkins etal. 1976; Perkins and
Turner 1988; Turner etal. 2001).
Notes: Neurospora is characterized by dark brown
ascospores with ornamented surface and its asexual morph
is usually found in nature and/or culture and is chrysonilia-
like (Shear and Dodge 1927; von Arx 1981a, b; Jacobson
etal. 2004). This genus is widespread and usually found in
soil and burned or unburned vegetation in terrestrial habitats
(Perkins etal. 1976; Perkins and Raju 1986; Perkins and
Turner 1988; Turner etal. 2001; García etal. 2004). Three
different mating strategies have been shown in Neurospora:
homothallism, pseudohomothallism and heterothallism
(Shear and Dodge 1927; Tai 1935; Mahoney etal. 1969;
Krug and Khan 1991; Krug etal. 1994). Initially, Gela-
sinospora was distinct from Neurospora as the latter has
longitudinal orientated structures on the ascospores, while
the former has foveolate ascospores (Dowding 1933). Phy-
logenetic analysis demonstrated that Gelasinospora and
Anixiella species scattered in Neurospora clades (García
etal. 2004). Hence, Gelasinospora and Anixiella were syn-
onymized under Neurospora (García etal. 2004; Kirk etal.
2008; Index Fungorum 2020). Pseudoneurospora was inde-
pendent from Neurospora because it has an irregular pat-
tern on the ascospores surface (García etal. 2004). Species
Fig. 41 Guilliermondia saccoboloides: a, b, e–l (PC-MNHN-PC-
PC0167691, holotype); c, d (redrawn from Boudier 1904). a Label
and herbarium material. b Ascomata. c Vertical view of ascoma. d
Ascus and paraphyses. e–h Asci (pedicel evanescent). i–l Ascospores.
Scale bars: b = 500µm, c = 200µm, d. h = 20µm, e–l = 10µm
Fungal Diversity
1 3
of Neurospora were traditionally distinguished based on
ascospore wall ornamentation (Dowding 1933; García etal.
2004; Cai etal. 2006b). Subsequently, García etal. (2004)
and Cai etal. (2006b) showed that the traditional classifica-
tion of Neurospora results in species being a polyphyletic in
the Sordariales in phylogenetic analysis. García etal. (2004)
considered the ornamentation of the epispore and the mating
strategy are the most important characters to consistent the
phylogenetic relatedness in Neurospora (Whitehouse 1949;
Taylor and Natvig 1989; Grognet and Silar 2015). In this
study, we were unable to find characteristics consistent with
polyphyletic Neurospora in phylogenetic analysis, either
ascospore morphology or mating strategies. Therefore, we
propose to delimit Neurospora as species nested in Sordari-
aceae in phylogenetic analysis and has aseptate ascospores
with various ribs and/or pitted on surface. In this study,
Neurospora is divided in four clades based on multi-gene
analysis. We have sorted out their main characteristics based
Fig. 42 Neurospora sitophila: a (redrawn from Shear and Dodge
1927); b–j (BPI-630404, from ex-type). a Conidiogenous cells
with conidia. b Culture. c Ascomata. d Vertical section of ascoma. e
Peridium. f, g Asci (evanescent). h–j Ascospores; Pseudoneurospora
amorphoporcata: k (redrawn from Udagawa 1980). k Ascospores.
Scale bars: a. h–k = 20µm, c = 1mm, d. e = 100µm, f. g = 50µm
Fungal Diversity
1 3
on mating type, form of ascomata, ascospore appearance and
types of epispore, and listed in Fig.40.
Type species: Neurospora sitophila Shear & B.O. Dodge, J.
Agric. Res., Washington 34: 1026 (1927)
Facesoffungi number: FoF 10024; Fig.42 a–j
Saprobic on wood and burnt vegetation, heterothallic.
Sexual morph: Ascomata 280–380µm (x̄ = 315µm, n = 10)
diam., perithecial, superficial, solitary, globose, dark brown
to black, with an ambiguous ostiole, covered by white hyphae.
Peridium 70–120µm (x̄ = 95µm, n = 30) wide, compr ising
four layers, outer layer composed of compact, dark brown
cells of textura angularis; middle layer closely aligned from
external to internal with hyaline to pale brown cells of textura
globulosa to textura angularis; and composed of brown cells
of textura angularis; inner layer composed of hyaline cells of
textura prismatica. Asci (150–)180–190(–200) × 13–16µm
(x̄ = 185 × 14.5µm, n = 30), 8-spored, unitunicate, cylindrical,
slightly curved, short pedicellate, thin-walled and evanescent
at maturity. Ascospores (17–)19–23(–28) × (10–)12–14(–15)
µm (x̄ = 22 × 13µm, n = 50), uni-seriate, subglobose to ellip-
soidal or oval, thick-walled, light brown when young, becom-
ing dark brown or olivaceous at maturity, aseptate, epispore
smooth, with a germ pore at each end. Ribs longitudinal or
reticulation, frequently branched, pigment deposited exter-
nally, nested straight, discrete, elongate pits. Asexual morph:
Hyphomycetous. Conidia orange, globose to subglobose,
aseptate, smooth-walled (adapted from García etal. 2004).
Material examined: Canada, Ottawa, on drying veneer
(birch and poplar) of fruit basket, culture 5574, 5 February
1927, M. Hantsport (BPI-630404, from ex-type).
Known hosts and distribution: On corn-meal agar cul-
ture of Monilia sitophila in Germany (type locality) (Shear
and Dodge 1927); in the air, on garbage, leaves, wood in
Australia, Canada, France, New Zealand and USA (Shaw
1998; Francuz etal. 2010; Vojtkova etal. 2020); on burnt
vegetation and non-burnt substrates distributed worldwide
(García etal. 2004).
Notes: Shear and Dodge (1927) found Neurospora sit-
ophila on a culture of Monilia sitophila and established
Neurospora. Subsequently, the hyphomycetous Chryson-
ilia sitophila and M. sitophila were synonymized under N.
sitophila (Perkins etal. 1976; Perkins and Turner 1988;
Turner etal. 2001; García etal. 2004), and this species was
found in the air and human waste, such as food or coffee
residues, and was identified as one of the causes of aller-
gies and asthma (Shaw 1998; Francuz etal. 2010; Vojtkova
etal. 2020). In this study, N. sitophila clustered in Neuros-
pora clade II accommodates heterothallic species (Fig.40).
Gelasinospora autosteira was considered as N. calospora
based on the similar morphology (García etal. 2004);
however, the ex-type strain of Gelasinospora autosteira
was sequenced by Vu etal. (2019), and this strain is sister
to N. nigeriensis in Neurospora clade III, whereas has far
distance with N. calospora. Therefore, we recommend
to transfer G. autosteira as Neurospora autosteira based
on phylogenetic analysis (see the combination list below,
Fig.40).
Neurospora tetraspora Dania García etal., in García etal.,
Mycol. Res. 108(10): 1134 (2004)
Basionym: Gelasinospora tetrasperma Dowding, Cana-
dian Journal of Research, Section C 9: 294 (1933)
Facesoffungi number: FoF 10025; Fig.43
Isolated from soil, dung or burnt vegetation, pseudoho-
mothallic. Sexual morph (On culture): Mycelium composed
of hyaline to light brown, branched, septate hyphae 3–5µm
wide (x̄ = 4µm, n = 30). Ascomata perithecial, solitary or
scattered, semi-immersed to superficial, slightly coriaceous,
pyriform, ostiolate, brown to black. Asci 4-spored, unitu-
nicate, with J-apical ring. Ascospores oval to ellipsoidal,
hyaline becoming dark brown, slightly flattened on one side,
aseptate, with a germ pore at each end, with circular pits.
Asexual morph: Undetermined (adapted from Dowding
1933; Matsushima 1975; von Arx 1982; Fort etal. 1990;
García etal. 2004).
Material examined: Culture from CBS, Netherlands (IMI-
38600, from ex-type).
Known hosts and distribution: On ptarmigan and rabbit
dung in Canada (type locality) (Dowding 1933); on dung and
soil in Canada, England, Finland, Norway, Russia, Spain,
Sweden and USA (Lundqvist 1972; García etal. 2004).
Notes: Dowding (1933) cultivated the spores which was
found on ptarmigan and rabbit dung in Manitobba and
Ontario and introduced a homothallic Gelasinospora tetrasp-
erma as the generic type of Gelasinospora. Dettman etal.
(2001) confirmed that G. tetrasperma is a pseudohomothal-
lic species. Subsequently, Gelasinospora was placed as a
synonym of Neurospora, and G. tetrasperma was replaced
by Neurospora tetraspora (García etal. 2004). Sequences
data from the ex-type strain (CBS:178.33) is available (Cai
etal. 2006b; Spatafora etal. 2006), and in this study, this
strain is distinct with N. tetraspora and nested in Neurospora
clade I which mainly comprises homothallic species with
an ornate epispore (Fig.40). We re-examined the ex-type
culture (38,600) of G. tetrasperma, and found that were no
ascomata, but a large number of conidia (22–28 × 12–15µm)
with a pitted surface in clumps on pale brown to brown,
septate, shrunken hyphae (Fig.43).
New combination:
Neurospora autosteira (Alexop. & S.H. Sung) S.K.
Huang & K.D. Hyde, comb. nov.
Basionym: Gelasinospora autosteira Alexop. & S.H.
Sung, Mycologia 42(6): 723 (1951) [1950]
Index Fungorum number: IF558325
Fungal Diversity
1 3
Pseudoneurospora Dania García etal., Mycol. Res. 108(10):
1139 (2004)
Isolated from soil. Sexual morph: Mycelium composed
of hyaline to brown, branched hyphae. Ascomata perithe-
cial or cleistothecial, gregarious or scattered, superficial to
immersed, yellowish to brown or black, sometimes translu-
cent, pyriform to subglobose, ostiolate, surrounded by hya-
line, septate hyphae or sinuose, brown, septate setae. Perid-
ium membranaceous, comprising brown to hyaline cells
of textura angularis or textura globulosa. Asci 8-spored,
unitunicate, cylindrical to clavate, short pedicellate, apex
rounded, with J- apical ring, evanescent. Ascospores uni- or
bi- seriate, hyaline when young becoming dark brown or
black, oval to ellipsoidal, aseptate, verrucose to tuberculate,
with a germ pore at each end. Asexual morph: Undeter-
mined (adapted from Udagawa 1980; García etal. 2004).
Notes: García etal. (2004) introduced Pseudoneurospora
for Gelasinospora amorphoporcata based on phylogenetic
analysis. They mentioned that this genus has ascospores
with a strongly verrucose to tuberculate irregular pattern
(Fig.42k) on the surface (Udagawa 1980; García etal.
2004). Subsequently, a second species, P. canariensis, was
introduced by Crous etal. (2014) having similar ascospore
morphology and molecular data. In this study, an invalid
name ‘Copromyces octosporus’ (CBS 386.78) is sister to P.
canariensis (see notes for Copromyces). Pseudoneurospora
species clustered with 92%ML/1.00PP support and basal to
Sordaria clade II in Sordariaceae (97%ML/1.00PP, Fig.40).
Type species: Pseudoneurospora amorphoporcata (Uda-
gawa) Dania García etal., Mycological Research 108 (10):
1139 (2004)
Basionym: Gelasinospora amorphoporcata Udagawa,
Transactions of the Mycological Society of Japan 21 (1):
19 (1980)
Facesoffungi number: FoF 10131; Fig.42k
Isolated from soil. Sexual morph: Ascomata perithe-
cial, brown to black, gregarious or scattered, superficial to
Fig. 43 Neurospora tetraspora:
(IMI-38600, from ex-type). a
Herbarium label. b Scattered
group of spores on mycelium. c
Group of conidia. d–h Conidia
(f showing pits, h showing
guttules). Scale bars: b = 1mm,
c = 200µm, d = 20µm,
e–h = 10µm
Fungal Diversity
1 3
immersed, pyriform to subglobose, membranaceous, ostio-
late, with papilla. Peridium comprising brown to hyaline
cells of textura angularis or textura globulosa, membrana-
ceous. Asci 8-spored, unitunicate, cylindrical, short pedicel-
late, apex rounded, with J- apical ring. Ascospores hyaline
when young becoming dark brown or black, oval to ellipsoi-
dal, aseptate, with a germ pore at each end, with irregular
reticulate patterns. Asexual morph: Undetermined (adapted
from Udagawa 1980; García etal. 2004).
Known hosts and distribution: On soil in Philippines and
Thailand (type locality) (Udagawa 1980; García etal. 2004).
Notes: Pseudoneurospora amorphoporcata was consid-
ered similar to Copromyces bisporus and Emblemospora
monotrema in having ornamented ascospores (García etal.
2004). However, Pseudoneurospora can be distinguished
from C. bisporus because the latter lacks an apical ring in
the asci and has globose ascospores with one germ pore.
It also can be differentiated from E. monotrema which has
ascospores with plate-like wrinkles (Lundqvist 1972; Jeng
and Krug 1976). In this study, P. amorphoporcata is basal
to P. canariensis and ‘Copromyces octosporus’ (91%ML,
Fig.40).
Sordaria Ces. & De Not., Comm. Soc. crittog. Ital. 1(4):
225 (1863)
Isolated from dung and soil or saprobic on wood. Sexual
morph: Ascomata perithecial, gregarious, immersed to
superficial, brown, translucent, subglobose to pyriform, gla-
brous or with rare hairs, ostiolate, with papilla, periphysate.
Peridium membranaceous, composed of dark brown to hya-
line cells of textura angularis to textura prismatica. Asci
8-spored, unitunicate, cylindrical, short pedicellate, apex
rounded, with J-apical ring, with or lacking paraphyses.
Ascospores uni-seriate, subglobose to ellipsoidal, hyaline,
becoming dark brown, aseptate, smooth-walled, germ pore
at one end, with or without a gelatinous sheath. Asexual
morph: Undetermined (adapted from De Notaris 1867;
Winter 1884; Potebnia 1907; Cain 1957a; Lundqvist 1972;
Khan and Krug 1989; Watanabe 1989).
Notes: Sordaria was first mentioned in Cesati and de
Notaris (1863), and typified by Sphearia sordaria Fr.,
which later was transferred as Coniochaeta sordaria
(Petrak 1953). Moreau (1953) introduced Sordaria fimicola
as its generic type, which has been widely accepted (von
Arx and Müller 1954; Munk 1957; Stafleu and Voss 1969;
Lundqvist 1972). Lundqvist (1972) clarified Sordaria spe-
cies as having dark perithecial ascomata, cylindrical asci
and brown, smooth-walled ascospores with or without a
gelatinous sheath. Traditionally, species of Sordaria are
coprophilous and usually found in faeces and soil (Kirk
etal. 2008; Maharachchikumbura etal. 2016). von Arx
etal. (1987) re-defined species with a gelatinous sheath
as Sordaria, while these lacking a gelatinous sheath were
placed in Asordaria. Subsequently, Cai etal. (2006b) pro-
posed that the presence or absence of the ascospore sheath
is not a phylogenetically significant for the separation of
Asordaria from Sordaria. We placed Asordaria under
Sordaria, and Sordaria is divided in three clades (Sord-
aria clade I, II, III) in Sordariaceae based on phylogenetic
analysis (Fig.40).
Type species: Sordaria fimicola (Roberge ex Desm.) Ces. &
De Not., Comm. Soc. crittog. Ital. 1(4): 226 (1863)
Basionym: Sphaeria fimicola Roberge ex Desm., Annls
Sci. Nat., Bot., sér. 3 11(2): 339 (1849)
Facesoffungi number: FoF 01150; Fig.44
Saprobic on dead wood, homothallic. Sexual morph:
Ascomata 250–350 × 190–300 µm (x̄ = 270 × 240 µm,
n = 10), perithecial, solitary or gregarious, immersed,
erumpent through bark of the host at maturity, globose to
subglobose, membranaceous, dark brown to black, ostiolate,
with papilla. Peridium 15–30µm wide, comprising two lay-
ers, outer layer composed of brown to dark brown cells of
textura angularis; inner layer composed of pale brown to
hyaline cells of textura prismatica. Paraphyses evanescent.
Asci (155–)165–180(–220) × 12–20µm (x̄ = 175 × 15µm,
n = 50), 8-spored, unitunicate, cylindrical to subclavate, short
pedicellate, apex truncate to rounded, with J- inconspicu-
ous apical ring. Ascospores (17–)19–23 × (9–)10–13 µm
(x̄ = 20 × 11.5µm, n = 50), uni-seriate, subglobose to oval or
ellipsoidal, smooth-walled, pale brown to dark brown, asep-
tate, germ pore at one end, with gelatinous sheath. Asexual
morph: Undetermined.
Material examined: Germany, Berlin, Brandenburg,
Berlin Bot. Garden, on rotten stems of Phragmites sp. with
Claviceps sp., May 1889, P. Hennings (S-F6838, lectotype
of Sordaria fimicola var. phragmiticola Henn.); Germany,
Hessen, Nassau, Östrich, on horse dung, 1802, L. Fuckel
(S-F6841, lectotype of Sphaeria equina Fuckel); Italy, on
decaying Vitis sp., 22 March 1930, Shear (BPI-580866).
Known hosts and distribution: On horse dung in France
(type locality) (Desmazières 1849); on hare, horse, guinea-
pig, camel and rabbit dung, heather soil, grass culms, fil-
ter paper, ergot-infested panicles of Phragmites communis
in Argentina, Austria, Belgium, Bulgaria, Canada, Czech
Republic, Denmark, England, Egypt, Finland, France, Ger-
many, Hungary, Italy, Netherlands, Norway, Poland, South
Africa, Spain and Sweden (Lundqvist 1972).
Notes: We re-examined three specimens, the lectotype
of Sordaria fimicola var. phragmiticola Henn. (F6838),
lectotype of Sphaeria equina Fuckel (F6841) and Sordaria
fimicola (BPI-580866). These authentic materials were
determined as Sordaria fimicola by Lundqvist (1972). Phy-
logenetically, the strains of Sordaria fimicola, which were
collected from Canada and Netherlands (Vu etal. 2019),
clustered in Sordaria clade II (62%ML, Fig.40).
Fungal Diversity
1 3
Strattoniaceae S.K. Huang, Maharachch. & K.D. Hyde,
fam. nov.
Facesoffungi number: FoF 10143; Index Fungorum num-
ber: IF558346
Etymology: Named for Strattonia, the type genus of this
family.
Coprophilous. Sexual morph: Ascomata perithecial, soli-
tary, scattered or gregarious, superficial to immersed, coria-
ceous to membranaceous, pyriform to subglobose, brown
to black, hairy, ostiolate, with a black papilla, periphysate.
Paraphyses filiform-ventricose, septate. Asci 4–8-spored,
unitunicate, cylindrical, apex truncate or rounded, with
J-apical ring. Ascospores uni-seriate, fusiform to obovoid,
hyaline to subhyaline, 1-septate near the base, sometimes
finely guttulae; upper cell becoming brown to black, swollen,
ellipsoidal, with an apical germ pore; pedicel short, conical
or cylindrical, hyaline, collapsing when dry, surrounded by
gelatinous sheath. Gelatinous sheath rounded or irregular,
swelling in water, invaginated at the pedicel, with indenta-
tions. Asexual morph: Hyphomycetous, phialophora-like.
Conidiophores arising from aerial hyphae, erect, inflated at
the base. Phialides monophialidic, smooth-walled, hyaline,
Fig. 44 Sordaria fimicola:
a, e, g, m–o (S-F6841); c, k
(S-F6838); b, d, f, h–j, l (BPI-
580866). a–c Ascomata on vari-
ous hosts. d Peridium crushed.
e Vertical section of ascoma.
f Peridium. g–j Immature to
mature asci (h in Melzer’s
reagent, j in cotton blue). k–o
Ascospores (m–o in Indian
ink). Scale bars: a. b = 500µm,
c = 200µm, e = 100µm,
g–j = 50µm, d. f = 20µm,
k–o = 10µm
Fungal Diversity
1 3
cylindrical, unbranched or branched. Conidia hyaline to pale
yellow, ellipsoidal to oval, aseptate, smooth-walled (adapted
from Lundqvist 1972; Bell 2000, 2005).
Type genus: Strattonia Cif. 1954
Notes: The coprophilous Strattonia was introduced based
on Fimetaria tetraspora and was initially characterized by
superficial to immersed, coriaceous ascomata, 4-spored
asci and brown ascospores (Ciferri 1954). Subsequently,
Lundqvist (1972) delimited this genus as having ostiolate
ascomata, non-amyloid asci and two-celled ascospores (a
dark brown fusiform or ellipsoidal upper cell and a hyaline
pedicel), sometimes with gelatinous sheath, which is simi-
lar to Podospora, Sordaria and Triangularia. Since authen-
tic material of S. tetraspora is lacking, Lundqvist (1972)
described this type species based on previous literature and
found a special feature, that is, if gelatinous sheath present, it
surrounds the ascospores but collapses at the pedicel. Eleven
species accepted in this genus based on morphology (Ciferri
1954; Lundqvist 1972; Abdullah 1983; Bell 2000, 2005).
Vu etal. (2019) sequenced Podospora petrogale and
Strattonia oblecythiformis. These two strains clustered
in 100%ML/1.00PP support as an unstable sister clade to
Diplogelasinospora (Fig.26). Morphologically, these two
species are coprophilous and have similar hairy ascomata,
non-amyloid, cylindrical asci and ascospores comprise a
fusiform to ellipsoidal, brown to dark brown upper cell and
a hyaline pedicel, while surrounded by gelatinous sheath,
except pedicel, in addition, both of them have phialophora-
like asexual morph (Bell 2000, 2005). These characters
are also similar to S. tetraspora. The difference between P.
petrogale and S. oblecythiformis is that the former has a
cylindrical pedicel in ascospores, whereas the latter has obl-
ecythiform pedicel (Bell 2000, 2005). Therefore, we placed
Podospora petrogale in Strattonia (see the combination list
below).
Strattonia carbonaria, S. karachiensis and S. minor with
no gelatinous sheath were placed in Jugulospora and Lun-
dqvistomyces (Neoschizotheciaceae) in phylogenetic analy-
sis (Marin-Felix etal. 2020; this study). Thus, we believe
that the gelatinous sheath surrounds the ascospores except
pedicel should be accepted as a feature for the identifying
Strattonia species.
We also raised Strattonia to a family level, Strattoniaceae.
It is characterized by two-celled ascospores comprise a
brown upper cell and a hyaline pedicel and surrounded by
a gelatinous sheath except for pedicel, which is different
from other lasiosphaeriaceous taxa. Phylogenetically, Strat-
tonia species form a sister clade to Diplogelasinosporaceae
(Fig.26).
Strattonia Cif., Sydowia 8(1-6): 245 (1954)
Coprophilous. Sexual morph: Ascomata perithecial,
solitary, scattered or gregarious, superficial to immersed,
coriaceous to membranaceous, pyriform to subglobose,
brown to black, hairy, ostiolate, with a black papilla, peri-
physate. Paraphyses filiform-ventricose, septate. Asci
4–8-spored, unitunicate, cylindrical, apex truncate or
rounded, with J- apical ring. Ascospores uni-seriate, fusi-
form to obovoid, hyaline to subhyaline, 1-septate near
the base, sometimes finely guttulae; upper cell becoming
brown to black, swollen, ellipsoidal, with an apical germ
pore; pedicel short, conical or cylindrical, hyaline, collaps-
ing when dry, surrounded by gelatinous sheath. Gelatinous
sheath rounded or irregular, swelling in water, invaginated
at the pedicel, with indentations. Asexual morph: Hypho-
mycetous, phialophora-like. Conidiophores arising from
aerial hyphae, erect, inflated at the base. Phialides mono-
phialidic, smooth-walled, hyaline, cylindrical, unbranched or
branched. Conidia hyaline to pale yellow, ellipsoidal to oval,
aseptate, smooth-walled (adapted from Lundqvist 1972; Bell
2000, 2005).
Notes: Strattonia is typified by S. tetraspora, which has
hairy ascomata, cylindrical asci and ascospores comprise
a brown upper cell and a hyaline pedicel and surrounded
by a gelatinous sheath except pedicel (Ciferri 1954; Lun-
dqvist 1972). Lundqvist (1972) proposed that Strattonia is
connected with Podospora, Sordaria and Triangularia and
tried to sort out the affinities of Strattonia species by mor-
phology. Subsequently, the no gelatinous sheath Strattonia
carbonaria, S. karachiensis and S. minor were placed in
Jugulospora and Lundqvistomyces respectively, based on
phylogenetic analysis (Miller and Huhndorf 2005; Vu etal.
2019, Marin-Felix etal. 2020; this study, see notes for Jugu-
lospora minor). We divide the remaining Strattonia species
that do not have receivable molecular data into three groups
according to the characteristics of their gelatinous sheath: (i)
Strattonia grandis, S. tetraspora and S. zopfii are similar to
S. petrogale and S. oblecythiformis have a gelatinous sheath
surrounds ascospores except pedicel (Lundqvist 1972; Bell
2005); (ii) Strattonia borealis and S. mesopotamica have
terminal gelatinous appendages surrounding apical germ
pore and pedicel (Lundqvist 1972; Abdullah 1983), which
are similar to Podospora fibrinocaudata in Bombardiaceae;
(iii) Strattonia dissimilis and S. insignis have ascospores sur-
rounded by a gelatinous sheath except apical germ pore and
pedicel (Lundqvist 1972; Barrasa etal. 1985), which are
similar to Apodospora gotlandica in Bombardiaceae. How-
ever, more fresh collections and molecular information of
Strattonia are expected to define its taxonomic placement
more accurately.
Type species: Strattonia tetraspora (R. Stratton) Cif.,
Sydowia 8(1-6): 245 (1954)
Basionym: Fimetaria tetraspora R. Stratton, Ohio St.
Univ. Bull. 26: 84 (1921)
Facesoffungi number: FoF 10132.
Fungal Diversity
1 3
On rotten paper. Sexual morph Ascomata perithecial,
solitary to gregarious, superficial to immersed, coriaceous,
pyriform to subglobose, black, ostiolate. Asci 4-spored,
unitunicate, cylindrical. Ascospores uni-seriate, fusiform to
obovoid, composed of a brown upper cell, with an apical
germ pore, and a hyaline pedicel, sometimes surrounded by
gelatinous sheath, but pedicel never. Asexual morph Unde-
termined (adapted from Ciferri 1954; Lundqvist 1972).
Known hosts and distribution: On rotten paper in Domi-
nica (type locality) (Ciferri 1954).
Notes: Ciferri (1954) introduced Strattonia tetraspora and
noted that this species was found on rotten paper from Moca,
Santiago, Dominican Republic, in April 1928. Lundqvist
(1972) mentioned that the authentic sample of S. tetras-
pora is lacking. In this study, we also could not obtain any
authentic specimens or illustrations of S. tetraspora. This
makes it impossible for us to identify its substrate, nor can
we provide more information about this species. However,
we provided the hand-drawing of Strattonia zopfii (Fig.45a),
which is similar to S. tetraspora has ascospores surrounded
by a gelatinous sheath but invaginated at the pedicel, refer
to Lundqvist (1972).
New combination:
Strattonia petrogale (A.E. Bell) S.K. Huang & K.D. Hyde,
comb. nov.
Basionym: Podospora petrogale A.E. Bell, Muelleria
12(2): 236 (2000) [1999]
Index Fungorum number: IF558343
Fig. 45 Strattonia zopfii: a (redrawn from Lundqvist 1972). a
Immature to mature ascospores surrounded by a gelatinous sheath
but invaginated at the pedicel; Jugulospora minor: b–j (UPS-
UPS:BOT:F-117927, holotype). b Herbarium material. c Asco-
mata on host. d Ascomata e Ascoma in cross section. f Asci.
g–j Ascospore. Scale bars: a = 50 µm, d = 200 µm, e. f = 100 µm,
g–j = 10µm
Fungal Diversity
1 3
Zygospermellaceae S.K. Huang, Maharachch. & K.D. Hyde,
fam. nov.
Facesoffungi number: FoF 10144; Index Fungorum num-
ber: IF558347
Etymology: Named for Zygospermella, the type genus of
this family.
Coprophilous, entomogenous or on the rock. Sexual
morph: Ascomata perithecial, solitary, superficial to semi-
immersed, obpyriform to subglobose, dark brown to black,
verrucose, ostiolate. Peridium comprising carbonaceous to
membranaceous, brown to black cells of textura angula-
ris to textura prismatica. Paraphyses unbranched, broad,
cylindrical, ventricose, septate. Asci 8-spored, unituni-
cate, cylindrical to subclavate, pedicellate, apex rounded,
with J- apical ring. Ascospores uni- or bi-seriate, hyaline,
aseptate, fusiform to ellipsoidal when young, becoming
elongate-subfusiform, ends truncate, 1–3-septate, con-
stricted at the middle septum, dark brown when mature,
with an appendage and germ pore at each end. Append-
ages terminal, hyaline, cylindrical, tapering, smooth-
walled, ends rounded, with longitudinally striate. Asex-
ual morph: Hyphomycetous. Conidiophores arising from
aerial hyphae, erect, inflated at the base. Conidia small,
globose to oval, forming globose masses on the apex of
the phialides (adapted from Cain 1934, 1935; Lundqvist
1972; Zhang etal. 2017).
Type genus: Zygospermella Cain 1935
Notes: Zygospermella was initially introduced as Del-
itschia (Pleosporales) because of the similar hourglass-
shaped ascospores, and then redefined as Zygospermella
and placed in Lasiosphaeriaceae (Mouton 1897; Cain 1935;
Lundqvist 1969). Phylogenetically, Zygospermella and Lasio-
sphaeris form a clade sister to Neoschizotheciaceae (Kruys
etal. 2014; Marin-Felix etal. 2020). In this study, Lasio-
sphaeris, Zygospermella, Episternus and Ramophialophora
petraea clustered as a sister clade of Neoschizotheciaceae
in LSU sequence data (not shown). However, in multi-gene
analysis, Episternus onthophagi, Ramophialophora petraea
and Zygospermella insignis form an independent cluster
(83%ML/0.99PP) and nested in Sordariales as an orphan
clade (83%ML/1.00PP, Fig.26). In addition, the phialophora-
like hyphomycetous E. onthophagi is epizoic fungus and R.
petraea was collected on the rock in Karst cave (Zhang etal.
2017; Górz and Boroń 2018). Zygospermella has special
hourglass-shaped ascospores attached to terminal append-
ages with longitudinally striate, which is different from the
other members in Sordariales.
Episternus Górz & Boroń, Phytotaxa 376(1): 49 (2018)
Type species: Episternus onthophagi Górz & Boroń,
Phytotaxa 376(1): 49 (2018)
Facesoffungi number: FoF 10133
Known hosts and distribution: On legs of dung beetle
(Onthophagus verticicornis) in Poland (type locality) (Górz
and Boroń 2018).
Notes: Górz and Boroń (2018) found the hyphae arising
from the surface of a dung beetle and has 1-septate, ovoid
conidia. The monotypic Episternus was established as an
epizoic fungus in Lasiosphaeriaceae based on LSU-SSU-
ITS combined sequence data (Górz and Boroń 2018). In this
study, the ex-type strain of E. onthophagi (KRAM F 58223)
is basal to Zygospermella insignis and Ramophialophora
petraea in Zygospermellaceae (100%ML/1.00PP, Fig.26).
Zygospermella Cain, Mycologia 27(2): 227 (1935)
Coprophilous. Sexual morph: Ascomata perithecial,
solitary, superficial to semi-immersed, obpyriform to sub-
globose, dark brown to black, verrucose, ostiolate. Perid-
ium comprising carbonaceous to membranaceous, brown to
black cells of textura angularis to textura prismatica. Para-
physes unbranched, broad, cylindrical, ventricose, septate.
Asci 8-spored, unitunicate, cylindrical to subclavate, pedi-
cellate, apex rounded, with J-apical ring. Ascospores uni-
or bi-seriate, hyaline, aseptate, fusiform to ellipsoidal when
young, becoming hourglass-shaped to elongate-subfusiform,
ends truncate, usually 1-septate, constricted at the middle
septum, dark brown when mature, with a germ pore and
appendage at each end. Appendages hyaline, cylindrical,
tapering, smooth-walled, ends rounded, with longitudinally
striate. Asexual morph: Undetermined (adapted from Cain
1934, 1935; Lundqvist 1972).
Notes: Zygospermella was invalidly introduced as
Zygospermum (Cain 1934). Cain (1935) proposed that
Zygospermum should be corrected to Zygospermella and
accepted Zygospermella insignis and Z. setosa. Lundqvist
(1969, 1972) clarified that these species are conspecific, as
Z. insignis, and introduced another species, Zygospermella
striata. Therefore, two species are presently accepted for
Zygospermella, Z. insignis (type) and Z. striata. Subse-
quently, Kruys etal. (2014) sequenced Z. insignis, which
was collected on muskox dung in the USA, cattle dung in
France and cow dung in Sweden, and found these strains
clustered in Lasiosphaeriaceae (s. lat.). In this study, Z.
insignis is sister to Ramophialophora petraea in Zygosper-
mellaceae (100%ML/1.00PP, Fig.26).
Type species: Zygospermella insignis (Mouton) Cain, Myc-
ologia 27(2): 227 (1935)
Facesoffungi number: FoF 10027; Fig.46
Coprophilous. Sexual morph: Ascomata
650–900 × 250–500µm, perithecial, solitary, superficial
to semi-immersed, carbonaceous to membranaceous, pyri-
form to subglobose, dark brown to black, verrucose, sur-
rounded by setae, ostiolate, with black, carbonaceous necks
Fungal Diversity
1 3
180–250 × 100–150µm, inner cells with hyaline periphyses.
Setae 2.5–4.5µm wide, brown, aseptate, tapering towards the
acute apex. Peridium 25–40µm wide, outer layer composed
of carbonaceous, brown to black cells of textura intricata to
textura angularis; inner layer composed of membranaceous,
hyaline cells of textura angularis to textura prismatica. Par-
aphyses 2.5–4.5µm wide (x̄ = 3.5µm, n = 30), unbranched,
cylindrical, septate, tapering, slightly constricted at the septa.
Fig. 46 Zygospermella insignis: a–k, m–p (BPI-618905); l
(redrawn from Cain 1934). a Herbarium material. b Ascoma semi-
immersed on host. c Vertical section of the neck. d Peridium. e Setae.
f Asci with paraphyses. g Paraphyses. h, i Asci. j Apical ring of
ascus. k–p Ascospores. Notes: f. i. j from permanent mount. Scale
bars: b = 500 µm, f = 100 µm, c. d. h. i = 50µm, e. g. k–p = 20 µm,
j = 10µm
Fungal Diversity
1 3
Asci (150–)200–240(–270) × 15–25µm (x̄ = 220 × 20µm,
n = 20), 8-spored, unitunicate, cylindrical to subclavate,
pedicellate, apex rounded and thickened. Ascospores uni-
or bi-seriate, hyaline, aseptate, fusiform to ellipsoidal when
young, becoming hourglass-shaped to elongate-subfusiform,
ends truncate, 1-septate, sometimes 3-septae, constricted at
the middle septum 3–7µm wide, dark brown when mature,
(26–)30–40(–45) × 7–10µm (x̄ = 35 × 8 µm, n = 20), with
a germ pore and appendage at each end. Appendages
4.5–8µm wide, hyaline, cylindrical, tapering, ends rounded,
smooth-walled, with longitudinally striate. Asexual morph:
Undetermined.
Material examined: Canada, Ontario. Bear Island, Lake
Temagami, on cow dung, 21 November 1933, R.F. Cain
(BPI-618905).
Known hosts and distribution: On dung of cow, sheep
and horse in Belgium (type locality), Canada, England, Nor-
way, Scotland, Spain and Sweden (Mouton 1897; Lundqvist
1969).
Notes: We re-examined authentic material collected by
Cain and determined by Lundqvist (mentioned in the label
of material) since the type specimen was unavailable.
Other genera inSordariales
Chaetomiaceae G. Winter
Stellatospora Tad. Ito & Nakagiri, Mycoscience 35 (4):
413 (1994)
Isolated from rice field soil. Sexual morph: Mycelium
white, superficial, effuse. Ascomata cleistothecial, solitary
or gregarious, globose to subglobose, dark brown to black,
covered in hyaline hairs. Peridium composed of brown to
hyaline cells of textura angularis. Asci 3–8-spored, unitu-
nicate, obpyriform to oval, evanescent. Ascospores multi-
seriate, stellate to irregular with up to 9 short protuberances
when mature, aseptate, pale greenish brown to brown, with a
germ pore on one protuberance, guttulate. Asexual morph:
Undetermined (adapted from Ito and Nakagiri 1994).
Notes: The monotypic Stellatospora typified by S. ter-
ricola has stellate ascospores and was initially placed in
Sordariaceae (Ito and Nakagiri 1994). Subsequently, the
ex-type strain (CBS 811.95) is available (Wang etal. 2019a,
b; Vu etal. 2019), and nests in Chaetomiaceae (Wang etal.
2019a, b; this study, 100%ML/1.00PP, Fig.26). Therefore,
this genus belongs in Chaetomiaceae based on phylogenetic
analysis.
Type species: Stellatospora terricola Tad. Ito & Nakagiri,
Mycoscience 35(4): 413 (1994)
Facesoffungi number: FoF 10028; Fig.47
Isolated from rice field soil, colonies on oatmeal agar
(OA). Sexual morph: Mycelium white, superficial, effuse.
Ascomata 75–120µm diam., cleistothecial, solitary or gre-
garious, globose to subglobose, dark brown to black, covered
in hyaline hairs. Peridium 3–5µm wide, composed of pale
brown to hyaline cells of textura angularis. Asci 3–8-spored,
unitunicate, obpyriform to oval, evanescent. Ascospores
(9–)11–15(–17) µm (x̄ = 12µm, n = 50) diam., multi-seriate,
stellate to irregular with up to 9 short protuberances when
mature, aseptate, pale greenish brown to brown, with a germ
pore on one protuberance, guttulate. Asexual morph: Unde-
termined (adapted from Ito and Nakagiri 1994).
Material examined: Japan, Ikeda, Osaka, rice field soil,
22 May 1990, T. Ito (NBRC-IFO-H-12166, holotype).
Known hosts and distribution: Rice field soil, Japan (type
locality) (Ito and Nakagiri 1994).
Notes: We re-examined the type material and found that
there are star-like ascospores filling the ascomata, but no
complete asci. Stellatospora terricola (CBS 811.95) is
basal to Ovatospora and Trichocladium in Chaetomiaceae
(100%ML/1.00PP, Fig.26).
Neoschizotheciaceae S.K. Huang & K.D. Hyde, fam. nov.
Facesoffungi number: FoF 10033; Index Fungorum num-
ber: IF558385
Etymology: Named for Neoschizothecium, the type genus
of this family.
Coprophilous or saprobic on wood or isolated from soil.
Sexual morph: Ascomata perithecial, solitary to scattered
or gregarious, immersed to semi-immersed or superficial,
obpyriform or ovoid, membranaceous to coriaceous, brown
to black, sometimes semi-transparent, ostiolate, with long or
short necks, glabrous or surrounded by hairs, periphysate.
Peridium comprising yellowish to pale brown or dark brown
cells of textura angularis, textura globulosa or textura epi-
dermoidea. Paraphyses absent, or cylindrical, septate. Asci
4 to multi-spored, unitunicate, cylindrical to clavate, pedi-
cellate, evanescent. Ascospores uni- or bi-seriate, ellipsoidal
or triangular, usually 0–1-septate, hyaline to dark brown,
with germ pore at one or each end, usually with gelatinous
appendages at one or each end, rarely surrounded by gelati-
nous sheath. Asexual morph: Hyphomycetous. Conidia glo-
bose to cylindrical, hyaline to pale brown, smooth-walled,
ovate masses on the apex of the phialides (adapted from
Marin-Felix etal. 2020).
Type genus: Neoschizothecium S.K. Huang & K.D. Hyde
Notes: The Neoschizotheciaceae clade accommodates
Apodus, Cercophora, Echria, Immersiella, Jugulospora,
Lundqvistomyces, Neoschizothecium, Pseudoechria, Pseu-
doschizothecium, Rinaldiella and Zygopleurage based on
the molecular data of their generic types, and also several
Arnium, Apiosordaria, Podospora, Ramophialophora and
Zopfiella species (88%ML, Fig.26). These taxa are similar
to Bombardiaceae, Podosporaceae and Lasiosphaeriaceae in
having ellipsoidal ascospores, sometimes with appendages,
Fungal Diversity
1 3
Fig. 47 Stellatospora terricola: a, e–p (NBRC-IFO-H-12166, holo-
type); b–d (redrawn from Ito and Nakagiri 1994). a Dry culture.
b Ascoma in cross section. c Ascus. e Ascomata with mycelium. f
Squashed ascoma. d, g–p Ascospores. Scale bars: b = 50 µm, c.
g–p = 10µm, d = 5µm, e = 200µm, f = 100µm
Fungal Diversity
1 3
but are phylogenetically distinct from these three families
(Fig.26). Marin-Felix etal. (2020) introduced this clade as
Schizotheciaceae based on Schizothecium. However, Schizo-
thecium is a synonym of Podospora (Podosporaceae) based
on the type species S. fimicola, which nests in Podospora
in the phylogenetic analysis (Wang etal. 2019a, b; Index
Fungorum 2020; this study, Fig.26). We introduce Neoschi-
zotheciaceae as a new family in Sordariales with 72%ML
support (Fig.26), based on Neoschizothecium. This family
includes 11 genera and is characterized by ostiolate asco-
mata, cylindrical to clavate asci and ellipsoidal ascospores,
sometimes with long or short cylindrical or lash-like gelati-
nous appendages. Marin-Felix etal. (2020) introduced three
genera Lundqvistomyces, Pseudoechria and Pseudoschizo-
thecium which we do not discuss in our study.
Apodus Malloch & Cain, Can. J. Bot. 49(6): 872 (1971)
Saprobic on Oryza sp. or coprophilous. Sexual morph:
Mycelium hyaline to brown, septate, branched hyphae.
Ascomata cleistothecial, solitary, superficial, globose to
subglobose, brown to black, surrounded by hairs. Peridium
composed of brown to hyaline cells of textura angularis.
Asci 8-spored, unitunicate, clavate to cylindrical, blunt at the
apex, evanescent. Ascospores bi-seriate or irregular, subglo-
bose to ellipsoidal, 0–1-septate, pale brown to dark brown,
with a germ pore at the apex, sometimes with hyaline sheath.
Asexual morph: Undetermined (adapted from Malloch and
Cain 1971).
Notes: Apodus is typified by A. deciduus in having asco-
mata with long hairs and ellipsoidal ascospores with a germ
pore at the apex (Malloch and Cain 1971). Subsequently, A.
oryzae was collected from a sheath of Oryza sativa in Italy,
characterized by lacking an ostiole, but with dark brown
ascomata covered with light brown, flexuous, septate hairs,
clavate asci and ellipsoidal to broadly fusiform, 0–1-sep-
tate ascospores with an apical germ pore and sometimes
covered with a hyaline, gelatinous sheath (von Arx 1975).
Apodus deciduus and A. oryzae were recognized as conge-
neric because of their similar morphology, but they do not
cluster together in a phylogenetic analysis (Cai etal. 2006b).
In this study, strains of A. deciduus and A. oryzae nested in
Neoschizotheciaceae and Naviculisporaceae, respectively
(Fig.26). Therefore, we place Apodus in Neoschizotheci-
aceae based on phylogenetic placement of the generic type,
A. deciduus. Apodus oryzae belongs to Naviculisporaceae
but requires further data.
Type species: Apodus deciduus Malloch & Cain, Can. J.
Bot. 49(6): 872 (1971)
Facesoffungi number: FoF 10029; Fig.48
Coprophilous. Sexual morph: Mycelium hyaline to
brown, septate, branched hyphae 2–4.5µm wide. Asco-
mata 195–500µm (x̄ = 300µm, n = 5) diam., cleistothecial,
solitary, superficial, globose to subglobose, brown to
black, surrounded by long, septate, brown, flexuous,
tapering hairs. Peridium 15–30 µm wide, composed
of brown to hyaline cells of textura angularis. Asci
(45–)52–58(–60) × (12–)14–17µm (x̄ = 55 × 15µm, n = 20),
8-spored, unitunicate, clavate to cylindrical, with short pedi-
cel, apex blunt, with an indistinct apical ring, evanescent.
Ascospores 12–16 × 8–11.5µm (x̄ = 14 × 9.5µm, n = 20), bi-
seriate or irregular, subglobose to ellipsoid, usually aseptate,
often producing a single septum in culture, pale brown to
dark brown, sometimes collapsing at the middle, with a germ
pore at the apex. Asexual morph: Undetermined (adapted
from Malloch and Cain 1971).
Material examined: USA, California, San Mateo Co.,
Crystal Springs Reservoir, on dung of dusky-footed wood
rat in nest, 30 December 1969, D. Malloch (TRTC-045704,
holotype).
Known hosts and distribution: On rat dung in the USA
(type locality) (Malloch and Cain 1971).
Notes: We requested the holotype but the sample was too
dry to find any clear structure except ascomata. The features
of ascospores were found from a permanent mount. Cai etal.
(2006b) sequenced the ex-type strain Apodus deciduus (CBS
506.70) which is basal to Cercophora newfieldiana and Zop-
fiella tardifaciens in Neoschizotheciaceae (100%ML/0.96PP,
Fig.26).
Cercophora Fuckel, Jb. nassau. Ver. Naturk. 23-24: 244
(1870) [1869-70]
Saprobic on woody or herbaceous plants or coprophil-
ous. Sexual morph: Ascomata perithecial, scattered to
gregarious, superficial to semi-immersed, globose to sub-
globose, brown to black, glabrous or hairy, ostiolate, with
papilla. Peridium composed of pale brown to brown cells of
textura angularis. Asci 8-spored, unitunicate, cylindrical,
pedicellate, evanescent. Ascospores bi-seriate, straight or
slight curved, ellipsoidal to cylindrical, aseptate, hyaline,
with guttules and bipolar appendages when young, becom-
ing 2–multi-celled, upper cell(s) swelling to become fusi-
form, dark brown when mature; lower cell(s) hyaline when
mature, smooth-walled, with guttules, appendages disappear
with age. Asexual morph: Hyphomycetous. Conidiophores
micronematous, reduced to conidiogenous cells. Conidia
asepate, hyaline to brown, smooth-walled, globose to sub-
globose or usually with a truncate base and rounded apex
(adapted from Fuckel 1870; del Valle Catania etal. 2011).
Notes: Cercophora is characterized by membranaceous to
carbonaceous ascomata and ascospores with a swollen head
and hyaline appendage(s) (Fuckel 1870). Chrysoporium,
Cladorrhinum and Phialophora species have been pub-
lished as the asexual morphs of Cercophora (Udagawa and
Muroi 1979; von Arx 1981a; Ueda 1994; del Valle Catania
etal. 2011). In this study, we found that Cercophora species
Fungal Diversity
1 3
are scattered in Lasiosphaeriaceae, Naviculisporaceae,
Podosporaceae and Neoschizotheciaceae in Sordariales
(Fig.26). Molecular data for the generic type, Cercophora
mirabilis, has been sequenced and found that this species
grouped with hyphomycetous Ramophialophora globispora
in Neoschizotheciaceae (Miller and Huhndorf 2005; Marin-
Felix etal. 2020; this study, 100%ML/1.00PP, Fig.26).
Therefore, we place this genus in Neoschizotheciaceae.
Type species: Cercophora mirabilis Fuckel, Jb. nassau. Ver.
Naturk. 23-24: 245 (1870) [1869-70]
Facesoffungi number: FoF 10030; Fig.49
Saprobic on woody or herbaceous plants or coprophil-
ous. Sexual morph: Ascomata perithecial, scattered,
semi-immersed, globose to subglobose, brown to black,
surrounded by hairs, ostiolate, with papilla. Peridium
composed of pale brown cells of textura angularis. Asci
Fig. 48 Apodus deciduus: a–e,
h–m (TRTC-045704 holotype);
f, g (redrawn from Malloch and
Cain 1970). a Herbarium mate-
rial label. b Herbarium material.
c Squashed ascoma. d Ascoma
on host. e, f Ascoma with long
hairs. g, h Asci. i Mycelium.
j–m Ascospores. Scale bars:
c, i = 100µm, d = 200µm,
e. f = 500µm, g. h = 20µm,
j–m = 10µm
Fungal Diversity
1 3
8-spored, unitunicate, cylindrical, pedicellate, apex rounded,
with subapical globule, evanescent. Ascospores bi-seriate,
slight curved, ellipsoid, aseptate, hyaline, with guttules
and bipolar appendages when young, becoming 2-celled,
upper cell smaller than lower cell: the upper cell swelling
to become fusiform, truncate at the base, dark brown when
mature, (15–)18–25 × 10–15µm (x̄ = 20 × 12µm, n = 20);
lower cell cylindrical, hyaline when mature, 32–40 × 4–5µm
(x̄ = 35 × 4.5 µm, n = 20), smooth-walled, with guttules,
appendages disappear with age. Asexual morph: Hypho-
mycetous. Conidiophores micronematous, reduced to con-
idiogenous cells. Conidia asepate, hyaline to brown, globose
to subglobose, smooth-walled (adapted from Udagawa and
Muroi 1979; del Valle Catania etal. 2011).
Material examined: Sri Lanka, Hambantota Distr., south-
ernmost part of Yala National Park near the coast, open dry
jungle, on buffalo (Bubalus bubalis) dung in moist chamber,
11 February 1974, N. Lundqvist (UPS-UPS:BOT:F-698102).
Known hosts and distribution: On rotten stalks of cabbage
in Germany (type locality) (Fuckel 1870); on dead woody or
herbaceous plants or dung in Brazil, Canada, Corsiea, Den-
mark, Finland, Germany, Italy, Morocco, Norway, Poland,
Scotland, Sri Lanka, Sweden (Lundqvist 1972).
Notes: Cercophora is typified by C. mirabilis (Fuckel
1870), which has vermiform ascospores with a swollen
head, hyaline appendage(s) and a phialophora-like species
was introduced as its asexual morph (Udagawa and Muroi
1979). Strain of C. mirabilis nested in Neoschizotheciaceae
(Miller and Huhndorf 2005; Marin-Felix etal. 2020). We
re-examined an authentic specimen determined by Lun-
dqvist (mentioned in the label of material). The type strain
Cercophora mirabilis (CBS 120402) is sister to Ramophi-
alophora globispora (LC6218) in Neoschizotheciaceae
(100%ML/1.00PP, Fig.26).
Echria (N. Lundq.) Kruys, Huhndorf & A.N. Mill., Fungal
Diversity 70: 106 (2014) [2015]
Coprophilous or saprobic on wood. Sexual morph: Asco-
mata perithecial, solitary, superficial to semi-immersed,
globose to subglobose, with rigid hairs, ostiolate, with
papilla. Asci 8-spored, unitunicate, cylindrical, pedicellate.
Ascospores uni-seriate, ellipsoidal to fusiform, aseptate,
smooth-walled or roughened wall, with a large central gut-
tule and surrounded by gelatinous sheath. Asexual morph:
Undetermined (adapted from Lundqvist 1972; Kruys etal.
2014).
Fig. 49 Cercophora mirabilis: a–e (UPS-UPS:BOT:F-698102); f–i (redrawn from Lundqvist 1972). a Herbarium material label. b Ascoma. c
Crushed peridium. d, e Ascospores with elongate pedicel. f Ascus. g, h Ascospores becoming maturity. Scale bars: b = 200µm, c–h = 50µm
Fungal Diversity
1 3
Notes: Echria initially was introduced as a subgenus of
Arnium and typified by Arnium macrotheca (Lundqvist
1972), and Kruys etal. (2014) raised Echria to generic level
and transferred two Arnium species, A. gigantospora and A.
macrotheca, to Echria based on LSU and TUB sequences
data. Subsequently, Marin-Felix etal. (2020) showed that
these two Echria species clustered and, in this study, two
Echria species clustered and is sister to Rinaldiella pentago-
nospora in Neoschizotheciaceae (100%ML/1.00PP, Fig.26).
Echria different from Rinaldiella in having subglobose asco-
mata with rigid hairs and ellipsoidal ascospores surrounded
by gelatinous sheaths, whereas Rinaldiella has subglobose
ascomata covered with hairs and clavate ascospores (Lun-
dqvist 1972; Crous etal. 2014).
Type species: Echria macrotheca (P. Crouan & H. Crouan)
Kruys, Huhndorf & A.N. Mill., Fungal Diversity 70: 106
(2014) [2015]
Basionym: Sphaeria macrotheca P. Crouan & H. Crouan,
Florule Finistère (Paris): 24 (1867)
Facesoffungi number: FoF 10134
Coprophilous or saprobic on wood. Sexual morph: Asco-
mata perithecial, solitary, superficial to semi-immersed, glo-
bose to subglobose, ostiolate, with papilla, surrounded by
brown setae. Asci 8-spored, unitunicate, cylindrical, pedi-
cellate. Ascospores uni-seriate, ellipsoidal to subglobose,
aseptate, with a large central guttule and surrounded by
gelatinous sheath. Asexual morph: Undetermined (adapted
from Lundqvist 1972; Kruys etal. 2014).
Known hosts and distribution: On old cow dung among
the Splachnum ampullaceum in France (type locality)
(Crouan and Crouan 1867); on dung of cow, hare, horse,
mule, rabbit, rat and sheep in Bulgaria, Canada, Denmark,
England, France, Morocco, Puerto Rico, Sweden and USA
(Lundqvist 1972).
Notes: Marin-Felix etal. (2020) established Schizothe-
ciaceae and placed E. macrotheca in this family based on
phylogenetic analysis. In this study, Echria macrotheca
is sister to E. gigantospora in Neoschizotheciaceae
(100%ML/1.00PP, Fig.26).
Immersiella A.N. Mill. & Huhndorf, Mycol. Res. 108(1):
31 (2004)
Saprobic on wood. Sexual morph: Ascomata perithecial,
solitary to scattered, immersed to semi-immersed, ovoid to
obpyriform, brown, verrucose, ostiolate, with papilla, covered
with brown hairs. Peridium comprising membranaceous, hya-
line to brown cells of textura angularis or textura prismatica.
Asci 8-spored, unitunicate, cylindrical, pedicellate, with J-
apical ring distinct, evanescent. Ascospores bi-seriate, cylin-
drical to geniculate, hyaline to pale brown, aseptate to multi-
septate, guttulate, smooth-walled, with gelatinous appendages
or appendage absent. Asexual morph: Undetermined (Miller
and Huhndorf 2004a; Kruys etal. 2014).
Notes: Miller and Huhndorf (2004a) established Immer-
siella with I. caudata and I. immersa (type) in Sordariales
based on phylogenetic analysis of LSU sequence data.
Subsequently, Kruys etal. (2014) found that the clade
comprising I. caudata and I. immersa was closely related
to Arnium hirtum, which was isolated from dung in Scot-
land. In our study, A. hirtum, I. caudata and I. immersa also
form a strongly supported clade in Neoschizotheciaceae
(100%ML/0.97PP, Fig.26). Therefore, we recommend to
treat A. hirtum as Immersiella hirta.
Type species: Immersiella immersa (P. Karst.) A.N. Mill.
& Huhndorf, Mycol. Res. 108(1): 31 (2004)
Basionym: Lasiosphaeria immersa P. Karst., Bidr. Känn.
Finl. Nat. Folk 23: 163 (1873)
Facesoffungi number: FoF 10031; Fig.50
Saprobic on wood. Sexual morph: Ascomata
270–450 × 360–460µm (x̄ = 325 × 370µm, n = 10), per i-
thecial, solitary to scattered, superficial to semi-immersed,
sitting on subiculum, ovoid to obpyriform, brown, membran-
aceous, verrucose, ostiolate, with papilla. Subiculum com-
posed of brown, septate, hyphae, 1.5–3.5µm wide. Peridium
55–95µm (x̄ = 75µm, n = 30) wide, comprising two layers,
outer layer composed of brown cells of textura angularis;
inner layer composed of hyaline cells of textura prismatica.
Paraphyses numerous, branched, septate, filiform. Asci
(200–)220–250(–270) × 14–17µm (x̄ = 235 × 15µm, n = 30),
8-spored, unitunicate, cylindrical, pedicellate, apex rounded,
with subapical globule, distinct J- apical ring, evanescent.
Ascospores 35–45(–50) × 4.5–6.5µm (x̄ = 42 × 5µm, n = 50),
bi-seriate, slight curved, cylindrical to geniculate, hyaline
and aseptate when young, becoming pale brown and multi-
septate, swelling at end when mature, with a large guttule
in each cell, smooth-walled, with a germ pore at each end.
Asexual morph: Undetermined.
Material examined: USA, Wisconsin, 45º 0.00ʹ N/90°
0.00ʹ W, on dead wood, 9 June 1999, S.M. Huhndorf
(F-SMH 4104).
Known hosts and distribution: On dead wood of Betula
in Finland (type locality) (Karsten 1873); on dead wood in
USA (Miller and Huhndorf 2004a).
Notes: Immersiella immersa is characterized by pyriform
ascomata with cylindrical to geniculate ascospores (Miller
and Huhndorf 2004a). We could not obtain the type material.
Therefore, we re-examined an authentic specimen collected
by Huhndorf.
New combination:
Immersiella hirta (E.C. Hansen) S.K. Huang & K.D.
Hyde, comb. nov.
Fungal Diversity
1 3
Basionym: Sordaria hirta E.C. Hansen, Vidensk. Meddel.
Dansk Naturhist. Foren. Kjøbenhavn 59: 22 (1877)
Synonym: Arnium hirtum (E.C. Hansen) N. Lundq. &
J.C. Krug, Symb. bot. upsal. 20(no. 1): 218 (1972)
Index Fungorum number: IF558326
Jugulospora N. Lundq., Symb. bot. upsal. 20(no. 1): 256
(1972)
Coprophilous. Sexual morph: Ascomata perithecial, gre-
garious, superficial to semi-immersed, ovoid to subglobose,
brown to black, membranaceous to carbonaceous, ostiolate,
with central papilla, periphysate. Peridium composed of
brown cells of textura angularis. Paraphyses numerous, fili-
form, flexuous, septate. Asci 8-spored, unitunicate, cylindri-
cal, pedicellate, apex truncate, with apical ring, evanescent.
Ascospores uni-seriate, broadly ellipsoidal to ovoid, with an
apical germ pore, hyaline when young, becoming upper cell
aseptate, brown to dark brown, with plate-like wrinkles on
surface when mature and truncate at the base; lower cell tiny,
hyaline conical. Asexual morph: Undetermined (adapted
from Lundqvist 1972).
Notes: Jugulospora is typified by J. rotula and char-
acterized by ovoid ascomata and ovoid ascospores with a
gelatinous cap-like cell at the base and a plate-like wrinkles
Fig. 50 Immersiella immersa: (F-SMH 4104). a Herbarium material
label. b, c Ascomata on host. d Ascoma in cross section. e Setae. f
Asci and paraphyses. g, h, n Asci. i–k Ascospores. l, m Germinated
ascospores. Scale bars: c = 500 µm, d = 200 µm, e. f = 100µm, g. h.
n = 50µm, i–m = 20µm
Fungal Diversity
1 3
(Lundqvist 1972). Marin-Felix etal. (2020) transferred this
genus to Schizotheciaceae and introduced three more spe-
cies, J. antarctrica, J. carbonaria and J. vestita based on
morphology and phylogenetic analysis. This study also con-
firms that Jugulospora strains nest in Neoschizotheciaceae
(56%ML/1.00PP, Fig.26).
Type species: Jugulospora rotula (Cooke) N. Lundq.,
Symb. bot. upsal. 20(no. 1): 260 (1972)
Basionym: Sphaeria rotula Cooke, Handb. Brit. Fungi
2: 868 (1871)
Facesoffungi number: FoF 10032; Fig.51
Fig. 51 Jugulospora rotula: (NY-01050505, Isolectotype). a Herbarium material. b Ascomata on host. c Ascoma in cross section. d Peridium.
e–i Asci (h. apical ring). j–q Ascospores. Scale bars: b = 200µm, c. e = 100µm, d. f. g. i = 50µm, j–q = 10µm
Fungal Diversity
1 3
Coprophilous. Sexual morph: Ascomata
230–250 × 150–230µm (x̄ = 240 × 170µm, n = 10), per i-
thecial, gregarious, superficial to semi-immersed, ovoid to
subglobose, brown to black, membranaceous to carbona-
ceous, ostiolate, with central, papilla, periphysate. Perid-
ium 15–45µm wide, composed of brown cells of textura
angularis. Paraphyses numerous, filiform, flexuous, sep-
tate. Asci (80–)100–120(–150) × 9–15µm (x̄ = 110 × 10µm,
n = 20), 8-spored, unitunicate, cylindrical, pedicellate,
apex truncate, with apical ring, evanescent. Ascospores
(10–)14–15(–16) × 8–10µm (x̄ = 14.5 × 9µm, n = 50), uni-
seriate, broadly ellipsoidal to ovoid, with an apical germ
pore, hyaline when young; becoming upper cell aseptate,
brown to dark brown when mature, with plate-like wrinkles
on surface and truncate at the base connected a tiny, hyaline
conical lower cell. Asexual morph: Undetermined.
Material examined: UK, Surrey, Abinger, on burnt ground,
October 1816, M.C. Cooke (NY-01050505, Isolectotype).
Known hosts and distribution: On burnt ground in Bel-
gium, Canada, Denmark and England (type locality) (Cooke
1871; Lundqvist 1972).
Notes: The sequence data of Jugulospora rotula were pro-
vided by Miller and Huhndorf (2005) and Marin-Felix etal.
(2020), and in this study, these strains form a sister clade to
J. minor (100%ML, Fig.26) in Neochizotheciaceae. We re-
examined a type of Jugulospora rotula which was collected
by Cooke (mentioned in the label of material).
New combination:
Jugulospora minor (N. Lundq.) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Strattonia minor N. Lundq., Symb. bot. upsal.
20(no. 1): 271 (1972)
Facesoffungi number: FoF 10,026; Index Fungorum num-
ber: IF558327; Fig.45 b–j
Isolated from soil. Sexual morph: Ascomata
300–400 × 320–340µm (x̄ = 350 × 330 µm, n = 5), perithe-
cial, solitary to gregarious, superficial to semi-immersed,
membranaceous, pyriform to subglobose, brown to black,
verrucose, ostiolate, with iso-radiating cells at surface,
inner cells with hyaline periphyses. Peridium 15–60µm
wide, outer layer composed of brown cells of textura
angularis; inner layer composed of hyaline cells of
textura prismatica. Paraphyses 6–10µm wide, filiform.
Asci (120–)160–170(–230) × 8–12 µm (x̄ = 165 × 10 µm,
n = 20), 8-spored, unitunicate, cylindrical. Ascospores
20–22(–25) × 6–10µm (x̄ = 21 × 8µm, n = 50), uni-seriate,
oval to ellipsoidal, with a germ pore at the apex, hyaline when
young, becoming upper cell aseptate, brown to dark brown
when mature, verrucose; lower cell small, hyaline, caudal,
triangular 2–3.5 × 2–3µm. Asexual morph: Undetermined.
Material examined: Sweden, Uppland, Uppsala,
Stadsskogen Forest, on burnt ground among Anthracobiae
and small mosses, N. Lundqvist (UPS-UPS:BOT:F-117927,
holotype).
Known hosts and distribution: On burnt soil in Belgium,
Denmark, France and Sweden (type locality) (Lundqvist
1972).
Notes: Strattonia is characterized by ascospores with a
large, brown cell and a short, hyaline cell, surrounded by a
gelatinous sheath (Lundqvist 1972). Miller and Huhndorf
(2005) and Vu etal. (2019) sequenced Strattonia carbonaria
and S. minor, and these two strains clustered with Jugulos-
pora in Neoschizotheciaceae (Fig.26). These two species,
have ascospores comprise a brown and a hyaline cell, but
gelatinous sheath absent (Lundqvist 1972), are more similar
to Jugulospora. Strattonia carbonaria has been transferred
as Jugulospora carbonaria (Marin-Felix etal. 2020). There-
fore, we transfer S. minor as Jugulospora minor based on
morphology and phylogenetic analyses.
Neoschizothecium S.K. Huang & K.D. Hyde, gen. nov.
Etymology: Name refers to genus similar to, but different
from Schizothecium
Facesoffungi number: FoF 10034; Index Fungorum num-
ber: IF558329
Saprobic on wood, coprophilous. Sexual morph: Asco-
mata perithecial, solitary, scattered or gregarious, superfi-
cial to semi-immersed, pyriform, olivaceous-brown to dark
brown, semi-transparent, surrounded by pale brown hyphae,
ostiolate, with papilla. Peridium comprising membrana-
ceous, pale brown cells of textura angularis. Paraphyses
cylindrical, septate. Asci 8–multi-spored, unitunicate, cylin-
drical, evanescent. Ascospores uni-seriate to irregularly
arranged, ellipsoidal to broadly fusiform, aseptate, hyaline to
dark brown, truncate at the base with a cylindrical, slender,
hyaline pedicel, umbonate apex, verrucose, with an apical
germ pore. Asexual morph: Undetermined (adapted from
Corda 1838; Lundqvist 1972; Munngai etal. 2012; Wang
etal. 2019a, b).
Notes: Schizothecium was introduced by Corda (1838)
and is typified by S. fimicola and has membranaceous asco-
mata and ascospores with lash-like caudae (Corda 1838;
Lundqvist 1972; Hu etal. 2006; Mungai etal. 2012). The
generic type, Schizothecium fimicola, was synonymized as
Podospora fimicola based on the epitype strain S. fimicola
(CBS 482.64), which nested in Podospora (Podosporaceae)
(Wang etal. 2019a, b; Index Fungorum 2020). Marin-Felix
etal. (2020) established Schizotheciaceae and raised Schi-
zothecium as its type genus; however, the generic type of
Schizothecium has not been clarified. Currently, Schizo-
thecium is a synonym of Podospora under Podosporaceae,
whereas several Schizothecium species clustered in
Schizotheciaceae (Marin-Felix etal. 2020; this study,
100%ML/1.00PP, Fig.26). In this study, a novel genus
Neoschizothecium is therefore proposed to accommodate
Fungal Diversity
1 3
species in the Neoschizotheciaceae clade (Fig.26) and is
typified by Neo. curvisporum (see the combination list
below). This genus nests in Neoschizotheciaceae (95%ML)
and is distinct from Podospora in multi-gene analysis
(Fig.26).
Type species: Neoschizothecium curvisporum (Cain) S.K.
Huang & K.D. Hyde, comb. nov.
Basionym: Sordaria curvispora Cain, Canadian Journal
of Research, Section C 26: 492 (1948)
Synonym: Schizothecium curvisporum (Cain) N. Lundq.,
Symb. bot. upsal. 20(no. 1): 334 (1972)
Facesoffungi number: FoF 10136; Index Fungorum num-
ber: IF558333; Fig.52q–r
Saprobic on seeds. Sexual morph: Ascomata perithecial,
gregarious, superficial, pyriform, olivaceous brown to dark
Fig. 52 Neoschizothecium conicum: a, b, e, h–l, p (F124745); c,
d, f, g, m, n (S-F124742); o (redrawn from Fuckel 1870). a Her-
barium material. b, c Ascoma on host. d Ascoma in cross section. e,
f Peridium. g, h Wall structure around ostiole with periphyses. i–k
Asci. l–p Ascospores; Neoschizothecium curvisporum: q, r (redrawn
from Cain 1948). q Asci. r Ascospores. Scale bars: b. c = 500 µm,
d = 200µm, e–g. l–p = 20µm, h–k = 50µm, q. r = 10µm
Fungal Diversity
1 3
brown, semi-transparent, surrounded by pale brown hyphae,
ostiolate, with papilla, periphysate. Peridium composed of
membranaceous, pale brown cells of textura angularis.
Asci 8-spored, unitunicate, cylindrical, apex rounded, pedi-
cellate. Ascospores uni- or bi-seriate, reniform, aseptate,
brown to dark brown, truncate at the base with a clavate,
hyaline pedicel, with an apical germ pore. Asexual morph:
Undetermined.
Known hosts and distribution: On seeds of Apium gra-
veolens and Daucus carota in the USA (type locality) (Cain
1948).
Notes: Schizothecium curvisporum was established
based on Sordaria curvispora which was isolated in culture
from seed of Daucus carota L. var. sativa DC. in the USA
(DAOM 20504) (Cain 1948; Lundqvist 1972). Subsequently,
Vu etal. (2019) sequenced the ex-type strain S. curvispo-
rum (CBS 506.50), and this strain nests in Neoschizothecium
(88%ML, Fig.26). In this study, we were unable to obtain
authentic materials. Ascomata of N. curvisporum is similar
to N. conicum and the hand-drawing is provided for the asci
and ascospores (Fig.52q–r) based on Cain (1948).
New combinations:
Neoschizothecium aloides (Fuckel) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Sordaria aloides Fuckel, Fungi rhenani exsic.,
suppl., fasc.: no. 2549 (1868)
Synonym: Schizothecium aloides (Fuckel) N. Lundq.,
Symb. bot. upsal. 20(no. 1): 253 (1972)
Index Fungorum number: IF558331
Neoschizothecium carpinicola (Mouch.) S.K. Huang &
K.D. Hyde, comb. nov.
Basionym: Podospora carpinicola Mouch., Persoonia
13(1): 197 (1986)
Synonym: Schizothecium carpinicola (Mouch.) L. Cai,
Fungal Diversity 19: 14 (2005)
Index Fungorum number: IF558332
Neoschizothecium conicum (Fuckel) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Cercophora conica Fuckel, Jb. nassau. Ver.
Naturk. 23-24: 245 (1870)
Synonym: Schizothecium conicum (Fuckel) N. Lundq.,
Symb. bot. upsal. 20(no. 1): 253 (1972)
Facesoffungi number: FoF 10035; Index Fungorum num-
ber: IF558328; Fig.52a–p
Coprophilous. Sexual morph: Ascomata
450–600 × 200–400µm (x̄ = 500 × 320µm, n = 5), perithe-
cial, solitary or gregarious, superficial to semi-immersed,
pyriform, brown to dark brown, membranaceous, ostiole,
central, papillate with iso-radiating cells, surrounded by
pale brown hyphae 2–4µm wide, with hyaline, septate
periphyses. Peridium 20–40µm (x̄ = 30µm, n = 30) wide,
comprising two layers, outer layer composed of pale brown
cells of textura angularis; inner layer composed of hyaline
cells of textura prismatica. Paraphyses tapering, cylindri-
cal, septate. Asci (150–)165–175(–195) × 15–25(–35) µm
(x̄ = 170 × 20µm, n = 30), 8-spored, unitunicate, cylindrical,
apex rounded, with thickened wall, pedicellate, evanescent.
Ascospores (16–)18–23(–25) × 10–16µm (x̄ = 20 × 13µm,
n = 50), uni-seriate, ellipsoidal to broadly fusiform, aseptate,
hyaline to dark brown, truncate at the base with a conical,
hyaline pedicel, umbonate apex with a cylindrical, evanes-
cent gelatinous appendage, verrucose, with an apical germ
pore, collapsing when dry. Asexual morph: Undetermined.
Material examined: Sweden, Öland, Ås par., Ottenby
lund., on fallen deer dung (Dama dama) in moist chamber in
Stockholm, 16 May 1984, N. Lundqvist (S-F124742); Swe-
den, Skåne, Östra Vemmerlöv par., W of Hörte., in seashore
meadow, on horse dung in moist chamber in Storvreta, 19
September 1996, N. Lundqvist (S-F124745).
Known hosts and distribution: On various kinds of dung,
particularly cow and horse dung in all over the Nordic area,
like Germany (type locality) and Sweden (Fuckel 1870; Lun-
dqvist 1972); on roots in Canada (Vu etal. 2019).
Notes: Schizothecium conicum was found with Cer-
cophora mirabilis on rotten cow dung under an oak tree
(Fuckel 1870). Subsequently, Lundqvist (1972) clarified
the relevant materials to Sch. conicum and proposed Sor-
daria curvula (= Podospora curvula) is a synonym. Vu
etal. (2019) sequenced P. curvula (= Sch. conicum) (CBS
434.50), and this strain clustered with other Schizothe-
cium strains with 50%ML support in Neoschizotheciaceae
(Marin-Felix etal. 2020; this study, Fig.26). In this study,
we placed Sch. conicum in Neoschizothecium and re-exam-
ined an authentic specimen collected by Lundqvist.
Neoschizothecium fimbriatum (A. Bayer) S.K. Huang &
K.D. Hyde, comb. nov.
Basionym: Sordaria fimbriata A. Bayer, Práce Mor.
Přirodověd. Společn. Brno 1(3): 83 (1924)
Synonym: Schizothecium fimbriatum (A. Bayer) Barrasa
& Soláns, Revta Ibér. Micol. 6(1): 3 (1989)
Index Fungorum number: IF558334
Neoschizothecium glutinans (Cain) S.K. Huang & K.D.
Hyde, comb. nov.
Basionym: Sordaria glutinans Cain, Univ. Toronto Stud.,
Biol. Ser. 38: 40 (1934)
Synonym: Schizothecium glutinans (Cain) N. Lundq.,
Symb. bot. upsal. 20(no. 1): 254 (1972)
Index Fungorum number: IF558338
Neoschizothecium inaequale (Cain) S.K. Huang & K.D.
Hyde, comb. nov.
Fungal Diversity
1 3
Basionym: Sordaria inaequalis Cain, Canadian Journal
of Research, Section C 26: 489 (1948)
Synonym: Schizothecium inaequale (Cain) N. Lundq.,
Symb. bot. upsal. 20(no. 1): 334 (1972)
Index Fungorum number: IF558339
Neoschizothecium minicaudum (Faurel & Locq.-Lin.) S.K.
Huang & K.D. Hyde, comb. nov.
Basionym: Podospora minicauda Faurel & Locq.-Lin.,
Revue Mycol., Paris 42: 344 (1978)
Index Fungorum number: IF558340
Neoschizothecium selenosporum (Stchigel, Guarro & M.
Calduch) S.K. Huang & K.D. Hyde, comb. nov.
Basionym: Podospora selenospora Stchigel, Guarro & M.
Calduch, Mycologia 94(3): 554 (2002)
Synonym: Schizothecium selenosporum (Stchigel, Guarro
& M. Calduch) Y. Marín & Stchigel, in Marin-Felix etal.,
Microorganisms 8(9, no. 1430): 34 (2020)
Index Fungorum number: IF558341
Neoschizothecium tetrasporum (G. Winter) S.K. Huang &
K.D. Hyde, comb. nov.
Basionym: Sordaria tetraspora G. Winter, Hedwigia 11:
161 (1872).
Synonym: Schizothecium tetrasporum (G. Winter) N.
Lundq., Symb. bot. upsal. 20(no. 1): 256 (1972)
Index Fungorum number: IF558342
Rinaldiella Deanna A. Sutton, Y. Marín, Guarro & E.H.
Thomps., in Crous etal., Persoonia 32: 301 (2014)
Human pathogen. Sexual morph: Ascomata perithecial,
immersed, scattered, pyriform to subglobose, membrana-
ceous, dark brown to black, surrounded by filiform hairs,
ostiolate, with papilla. Peridium membranaceous, translu-
cent, composed of brown to yellowish brown cells of textura
epidermoidea. Paraphyses filiform, septate. Asci 8-spored,
unitunicate, clavate to cylindrical, pedicellate, evanescent.
Ascospores uni- to bi-seriate, hyaline, clavate and aseptate
when young, upper cell becoming swollen to polygonal,
brown, with truncate base and slightly acuminate apex;
lower cell conical, subhyaline, slightly warted. Asexual
morph: Undetermined (adapted from Crous etal. 2014).
Notes: The monotypic genus Rinaldiella was isolated
from a contaminated human lesion and is characterized by
pyriform ascomata with filiform brown hairs and clavate,
1-septate ascospores with a swollen, verrucose upper cell
(Crous etal. 2014). It is different from Echria has subglo-
bose ascomata with rigid hairs and ellipsoidal ascospores
surrounded by a gelatinous sheath (Lundqvist 1972). In this
study, we place this species to Neoschizotheciaceae, based
on its ex-type strain, Rinaldiella pentagonospora (CBS
132344) which is sister to Echria in Neoschizotheciaceae
(100%ML/1.00PP, Fig.26).
Type species: Rinaldiella pentagonospora Deanna A. Sut-
ton, Y. Marín, Guarro & E.H. Thomps., in Crous etal., Per-
soonia 32: 301 (2014)
Facesoffungi number: FoF 10135; Fig.37h–i
Human pathogen. Sexual morph: Ascomata perithecial,
immersed, scattered, pyriform to subglobose, membrana-
ceous, dark brown to black, surrounded by filiform hairs,
ostiolate, with papilla. Peridium membranaceous, translu-
cent, composed of brown to yellowish brown cells of textura
epidermoidea. Paraphyses filiform, septate. Asci 8-spored,
unitunicate, clavate to cylindrical, pedicellate, evanescent.
Ascospores uni- to bi-seriate, hyaline, clavate and aseptate
when young, upper cell becoming swollen to polygonal,
brown, with truncate base and slightly acuminate apex;
lower cell conical, subhyaline, slightly warted. Asexual
morph: Undetermined (adapted from Crous etal. 2014).
Known hosts and distribution: On human lesion in the
USA (type locality) (Crous etal. 2014).
Notes: Rinaldiella is sister to Echria (100%ML/1.00PP,
Fig.26). The former is characterized by ascospores com-
prise a brown, polygonal upper cell and a subhyaline, conical
lower cell, but different from the latter in having ellipsoidal
ascospores surrounded by a gelatinous sheath (Crous etal.
2014; Kruys etal. 2014).
Zygopleurage Boedijn, Persoonia 2(3): 316 (1962)
Coprophilous. Sexual morph: Ascomata perithecial, soli-
tary to scattered, superficial to semi-immersed, semi-trans-
parent, pyriform to subglobose, olivaceous-brown to black,
verrucose, ostiolate, with short necks. Peridium comprising
membranaceous, hyaline to brown cells of textura angu-
laris. Asci 4–8-spored, unitunicate, cylindrical to clavate,
pedicellate, apex rounded. Ascospores irregularly arranged
or spirally coiled around each other, hyaline, cylindrical
to fusiform, aseptate when young, becoming elongate, the
terminal cells swelling fusiform or oval and dark brown or
olivaceous-brown with truncate base; the intercalary cell
elongate, hyaline, sometimes slightly swelling when mature,
usually 3-celled, sometimes multi-septate, with gelatinous
appendages. Asexual morph: Undetermined (adapted from
Boedijn 1962; Mirza and Nasir 1968; Lundqvist 1969, 1972).
Notes: All three species of Zygopleurage were collected
from dung, especially cow dung, and the generic type, Zygo-
pleurage zygospora, was first found in Italy and Zygopleur-
age faiyumensis and Z. multicaudata were collected in Egypt
and Pakistan respectively (Boedijn 1962; Mirza and Nasir
1968; Lundqvist 1969). Lundqvist (1972) proposed that this
genus is similar to Lasiosphaeria and Podospora based on
ascospores comprise two brown cells connected by a long
Fungal Diversity
1 3
hyaline cell. Huhndorf etal. (2004b) sequenced Z. zygospora
(SMH4219). In this study, this strain nested in Neoschizo-
theciaceae with 94%ML support (Fig.26).
Type species: Zygopleurage zygospora (Speg.) Boedijn, Per-
soonia 2(3): 316 (1962)
Basionym: Sordaria zygospora Speg., Michelia 1(no. 2):
227 (1878).
Facesoffungi number: FoF 10036; Fig.53
Coprophilous. Sexual morph: Ascomata
0.7–1 × 0.5–0.8mm, perithecial, solitary to scattered, super-
ficial to semi-immersed, membranaceous, semi-transparent,
pyriform to subglobose, olivaceous-brown, verrucose, ostio-
late, with dark brown necks 300–400 × 150–300µm, inner
cells with hyaline periphyses. Peridium 25–60µm wide,
outer layer composed of brown cells of textura angularis;
Fig. 53 Zygopleurage zygospora: a–h (S-F43798); i (redrawn from Lundqvist 1969). a Herbarium material label. b Ascoma on host. c Ascoma
in cross section. d Neck with periphyses. e Peridium. f–h Asci. i Ascospores. Scale bars: b = 500µm, c = 200µm, d. f–h = 100µm, e. i = 50µm
Fungal Diversity
1 3
inner layer composed of hyaline cells of textura prismat-
ica. Asci (250–)280–315(–330) × (35–)40–50(–55) µm
(x̄ = 300 × 45µm, n = 20), 8-spored, unitunicate, cylindri-
cal to clavate, pedicellate, apex rounded and thick-walled.
Ascospores bi-seriate, 3-celled, the two terminal cells hya-
line to dark brown, ellipsoidal to fusiform with an apical
germ pore and truncate at the base, (25–)27–32(–35) × (12–
)14–18(–20) µm (x̄ = 30 × 15µm, n = 50), with 4 distinct hya-
line gelatinous processes at each end; connected by a long,
hyaline 4–7µm wide cell, with gelatinous ornamentation,
evanescent. Asexual morph: Undetermined.
Material examined: Egypt, Faiyum Province, Al Fayyum,
Libyan Desert, along the Cairo-Faiyum desert road, in the
dried-up salt marsh at Kom Aushim 10kmN.E. of Lake
Qarun, on cow dung in moist chamber in Uppsala, 1 March
1968, N. Lundqvist (S-F43798); Sweden, Småland, Vim-
merby parish, 3km S of Storebro, on cow dung after 3weeks
in moist chamber in Uppsala, 31 May 1959, N. Lundqvist
(S-F43801).
Known hosts and distribution: On dung of cow, guinea
pig, horse in Canada, Egypt, France, Italy (type locality),
Liberia, Puerto Rico, Sweden and USA (Boedijn 1962; Lun-
dqvist 1969).
Notes: We re-examined the authentic specimen collected
by Lundqvist (mentioned in the label of material).
Sordariales genera incertae sedis
Arnium Nitschke ex G. Winter, Bot. Ztg. 31: 450 (1873)
Saprobic on wood, coprophilous. Sexual morph: Asco-
mata perithecial, scattered, superficial to semi-immersed,
obpyriform to subglobose, membranaceous, pale brown to
dark brown, ostiolate, with necks. Necks cylindrical to coni-
cal, black, glabrous or covered with unbranched, tapering,
brown hairs, with hyaline, filiform periphyses. Paraphyses
filiform to cylindrical, septate. Asci 8-spored, unitunicate,
cylindrical to clavate, pedicellate, apex rounded, evanescent.
Ascospores uni-seriate to irregularly arranged, obovoid to
ellipsoidal to fusiform, aseptate, hyaline when young,
becoming dark brown when mature, with ornamented wall
and a gelatinous appendage at each end or surrounded by a
gelatinous sheath. Asexual morph: Undetermined (adapted
from Nitschke 1873; Lundqvist 1974).
Notes: Arnium is typified by A. lanuginosum and has sub-
globose ascomata, cylindrical to clavate asci and ascospores
with a gelatinous appendage at each end (Nitschke 1873).
Subsequently, A. lanuginosum was synonymized under
Podospora brassicae (Ellis and Everhart 1892). This genus
was rarely mentioned until it was clarified by Lundqvist
(1972), and several Arnium species were then introduced
(Cain and Mirza 1972; Krug and Cain 1972; Lundqvist
1974). Arnium species are phylogenetically scattered among
the families of Sordariales, and several taxa have been trans-
ferred to Cladorrhinum, Podospora and Triangularia (Wang
etal. 2019a, b; Marin-Felix etal. 2020). In this study,
Arnium species nested in Naviculisporaceae, Podosporaceae
and Neoschizotheciaceae, but there is no molecular data for
A. lanuginosum. Thus, we place the genus in Sordariales
genera incertae sedis pending more data.
Type species: Arnium lanuginosum Nitschke [as 'lanugi-
nosa'], Bot. Ztg. 31: 450 (1873)
Facesoffungi number: FoF 10137
Saprobic on wood. Sexual morph: Ascomata perithecial,
superficial, globose to subglobose, tomentose. Asci 8-spored,
unitunicate, cylindrical. Ascospores oval to ellipsoidal, dark
brown, with gelatinous appendage at each end. Asexual
morph: Undetermined (adapted from Nitschke 1873; Ellis
and Everhart 1892).
Notes: Ellis and Everhart (1892) believed that Arnium lanug-
inosum and Podospora brassicae were conspecific, and these
species were synonymized under A. olerum (Lundqvist 1972)
based on similar ellipsoidal ascospores with cylindrical gelati-
nous appendages. In this study, A. olerum was transferred as C.
olerum based on phylogenetic analysis (see notes for Clador-
rhinum olerum). However, there is no molecular data for A.
lanuginosum and we could not obtain any related specimens.
Biconiosporella Schaumann, Veröff. Inst. Meeresf. Bremer-
haven 14(1): 24 (1972)
Saprobic on wood. Sexual morph: Ascomata perithecial,
scattered, semi-immersed to superficial, globose to subglo-
bose, collapsing when dry, membranaceous, black, ostiolate,
with papilla. Asci 8-spored, unitunicate, cylindrical, pedicel-
late, apex rounded, evanescent. Ascospores uni-seriate, obo-
void to ellipsoidal, hyaline when young, becoming pale brown
to brown when mature, smooth-walled, 3-septate, with a germ
pore at each end and with tubercules at the middle. Asexual
morph: Undetermined (adapted from Schaumann 1972).
Notes: Biconiosporella is characterized by obovoid
to ellipsoidal ascospores with tubercules at the middle
(Schaumann 1972). Jones etal. (2009) referred this genus
to Lasiosphaeriaceae based on morphological observations.
We found that its unique ascospores are similar to Stellatos-
pora (Chaetomiaceae). Molecular data for B. corniculate is
lacking, therefore, we place Biconiosporella in Sordariales
genera incertae sedis pending fresh collection.
Type species: Biconiosporella corniculata Schaumann,
Veröff. Inst. Meeresf. Bremerhaven 14(1): 24 (1972)
Facesoffungi number: FoF 10037; Fig.54
Saprobic on wood. Sexual morph: Ascomata
275–470 × 150–240µm (x̄ = 370 × 180 µm, n = 5), peri-
thecial, scattered, semi-immersed to superficial, glo-
bose to subglobose, collapsing when dry, membrana-
ceous, black, ostiolate, with papilla. Peridium comprising
dark brown to hyaline cells of textura angularis. Asci
Fungal Diversity
1 3
(200–)220–250(–300) × 20–35 µm (x̄ = 230 × 26 µm,
n = 20), 8-spored, unitunicate, cylindrical, pedicellate, apex
rounded, evanescent. Ascospores 28–35(–38) × 19–25µm
(x̄ = 33 × 22.5µm, n = 50), uni-seriate, obovoid to ellipsoi-
dal, hyaline when young, becoming pale brown to brown
when mature, smooth-walled, with a germ pore at each end
and 3–4 obvious coniform tubercules evenly distributed at
middle, 3-septate, one septum near apex, two septa near the
truncate base. Asexual morph: Undetermined.
Material examined: Germany, on rotten wood in sea
water, 25 November 1969, K. Schaumann (NY-01388911,
holotype).
Known hosts and distribution: On dead wood in Germany
(type locality) (Schaumann 1972).
Notes: The monotypic Biconiosporella has unique
ascospores with coniform tubercules at center. We re-exam-
ined the holotype which is well preserved, but there were no
mature asci.
Fig. 54 Biconiosporella
corniculata: (NY-01388911,
holotype). a Herbarium mate-
rial label. b Scattered ascomata.
c Ascoma on host. d Col-
lapsing ascoma. e Squashed
ascoma. f Ostiole. g Apex of
ascus. h Ascus (immature).
i–l Ascospores. Scale bars:
c = 500µm, d. e = 200µm,
f = 100µm, h = 50µm,
g = 20µm, i–l = 10µm
Fungal Diversity
1 3
Camptosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23-24: 140
(1870) [1869-70]
Saprobic on stems or wood or coprophilous. Sexual
morph: Ascomata perithecial, scattered, superficial to
semi-immersed, erumpent through bark of host, pyriform
to subglobose, black, glabrous or hairy, ostiolate, with
papilla, periphysate. Peridium membranaceous. Paraphyses
filiform. Asci 8-spored, unitunicate, cylindrical to clavate,
pedicellate, with J- apical ring. Ascospores broadly clavate
to obpyriform, aseptate, hyaline to brown, smooth-walled or
verrucose, with conical gelatinous appendage at the apex.
Asexual morph: Undetermined (adapted from Fuckel 1870;
Lundqvist 1972).
Notes: Fuckel (1870) established Camptosphaeria which
is typified by Cam. sulphurea. Lundqvist (1972) reduced
this genus as a subgenus of Cercophora because the cylin-
drical to broadly fusiform ascospores are similar to Cer-
cophora and Podospora. Krug and Jeng (1977) re-defined
Camptosphaeria as a genus which has pod-like ascospores
with conical gelatinous appendages and suggested that its
ascospores are similar to Bombardia (Bombardiaceae)
and Lasiosphaeria (Lasiosphaeriaceae). Thus, we place
the genus in Sordariales genera incertae sedis pending
molecular data.
Type species: Camptosphaeria sulphurea Fuckel, Jb. nas-
sau. Ver. Naturk. 23-24: 140 (1870) [1869-70]
Facesoffungi number: FoF 10038; Fig.55
Saprobic on stems. Sexual morph: Ascomata
520–650 × 400–450µm (x̄ = 560 × 420µm, n = 5), perithe-
cial, scattered, semi-immersed, erumpent through bark of
host, pyriform to subglobose, black, ostiolate, with papilla,
periphysate. Peridium 20–50µm (x̄ = 30µm, n = 30) wide,
membranaceous, comprising two layers, outer layer com-
posed of brown cells of textura angularis; inner layer
composed of hyaline cells of textura prismatica. Para-
physes filiform. Asci (130–)140–170(–180) × 12–25 µm
(x̄ = 150 × 18µm, n = 30), 8-spored, unitunicate, cylindrical,
pedicellate, apex rounded, with apical globule. Ascospores
(25–)28–30(–35) × 10–15µm (x̄ = 30 × 13µm, n = 30), bi-
seriate, cylindrical to broadly clavate to obpyriform, asep-
tate, hyaline to pale brown, smooth-walled, with conical
gelatinous appendage at the apex. Asexual morph: Unde-
termined (adapted from Fuckel 1870; Lundqvist 1972).
Material examined: Germany, Hessen, Nessen, Östrich,
on stems of Peucedanum officinale, 1894, L. Fuckel
(G-318793/1, holotype).
Known hosts and distribution: On stems of Peucedanum
in Germany (type locality) (Fuckel 1870).
Notes: Fuckel (1870) introduced Camptosphaeria sul-
phurea with pod-like ascospores having conical gelatinous
appendages. We re-examined the holotype (318793/1) which
was determined by Lundqvist. Complete asci and ascospores
were not observed, so we drew them with reference to Fuckel
(1870) and Lundqvist (1972).
Diffractella Guarro, P.F. Cannon & Aa, Syst. Ascom. 10:
107 (1991)
Saprobic on wood. Sexual morph: Ascomata cleistothe-
cial, solitary, superficial, carbonaceous, globose, with alveo-
late-reticulate ornamentation, brown to black, surrounded by
long setae. Peridium composed of brown to reddish brown,
radial, septate, cylindrical cells. Asci 8-spored, unitunicate,
globose to oval. Ascospores multi-seriate, fusiform to falci-
form, upper cell(s) brown to dark brown, aseptate or with
1-septate near the base, guttulate; lower cell cap-like, hya-
line. Asexual morph: Undetermined (adapted from Fuckel
1872; Guarro etal. 1991).
Notes: Guarro etal. (1991) transferred Cephalotheca cur-
vata (Fuckel 1872) to Diffractella as the generic type, which
is characterized by cleistothecial ascomata, globose asci and
fusiform ascospores with cap-like basal cell. The genus was
considered as a member of Lasiosphaeriaceae because it
has falciform ascospores with a special cap-like cell, which
is similar to Bellojisia and Zopfiella (Guarro etal. 1991;
Réblová 2008).
Type species: Diffractella curvata (Fuckel) Guarro, P.F.
Cannon & Aa, Syst. Ascom., Reprint of Volumes 1-4 (1982-
1985) 10(2): 108 (1991)
Basionym: Cephalotheca curvata Fuckel, Jb. nassau. Ver.
Naturk. 25–26: 298 (1871)
Facesoffungi number: FoF 10039; Fig.56
Saprobic on wood. Sexual morph: Ascomata
245–400µm (x̄ = 300µm, n = 5) diam., cleistothecial, soli-
tary, superficial, carbonaceous, globose, with alveolate-
reticulate ornamentation, brown to black, surrounded by
long setae. Setae 4–6µm wide, filiform, septate, brown
and tapering, lighter at the rounded apex. Peridium com-
posed of brown to reddish brown, radial-shaped, sep-
tate, cylindrical cells. Asci 15–22µm (x̄ = 18 µm, n = 20)
diam., 8-spored, unitunicate, globose to oval. Ascospores
15–18(–21) × 4.5–7.5 µm (x̄ = 17 × 6µm, n = 20), multi-
seriate, fusiform to falciform, upper cell(s) brown to dark
brown, aseptate or with 1-septate near the base, guttulate;
lower cell cap-like, hyaline, sometimes collapsing when dry.
Asexual morph: Undetermined.
Material examined: Germany, Baden-Wurttemberg, Ca.
Eberbach, 9 º0′11’’E/48 º35′33’’N, on rotten oak wood,
1894, L. Fuckel (G-352276/1, holotype).
Known hosts and distribution: On rotten hollow trunks of
oak in Germany (type locality) (Fuckel 1872).
Notes: We re-examined the holotype of Diffractella cur-
vata (G-352276/1).
Fungal Diversity
1 3
Emblemospora Jeng & J.C. Krug, Can. J. Bot. 54(16): 1971
(1976)
Coprophilous. Sexual morph: Ascomata perithecial,
solitary, superficial to semi-immersed, pyriform to subglo-
bose, brown to black, ostiolate, with black necks, surrounded
by setae. Peridium membranaceous, outer layer composed
of brown cells of textura angularis; inner layer composed
of hyaline cells of textura prismatica. Asci 8-spored,
unitunicate, subglobose to clavate, pedicellate, apex rounded
or truncate, with J-apical ring, evanescent. Ascospores bi-
seriate, ellipsoidal to subglobose, brown to dark brown,
aseptate, with plate-like wrinkled wall, germ pore at one
or both end(s), guttulate. Asexual morph: Undetermined
(adapted from Jeng and Krug 1976).
Notes: Jeng and Krug (1976) established Emblemospora,
which includes two species, E. ditrema and E. monotreme
Fig. 55 Camptosphaeria sulphurea: a–i (G-318793/1, holotype);
j–k (redrawn from Fuckel 1870 and Lundqvist 1972). a Herbarium
material label. b Herbarium material. c Ascomata on host. d Ascoma
in cross section. e Crushed papilla with periphyses. f Peridium. g–j
Asci. k Ascospores. Scale bars: d = 200µm, e = 100µm, f–k = 50µm
Fungal Diversity
1 3
(type), in Sordariaceae. Species are similar to Diplogelasi-
nospora, Jugulospora and Neurospora in having pyriform
ascomata and clavate asci with one-celled ascospores with
ornamentation (Jeng and Krug 1976). Molecular data is una-
vailable for this genus.
Type species: Emblemospora monotrema Jeng & J.C. Krug,
Can. J. Bot. 54(16): 1972 (1976)
Facesoffungi number: FoF 10040; Fig.57
Coprophilous. Sexual morph: Ascomata
450–600 × 230–380µm (x̄ = 500 × 300 µm, n = 5), peri-
thecial, solitary, superficial to semi-immersed, pyri-
form to subglobose, brown to black, membranaceous,
ostiolate, with black necks (65–100 × 100–120µm), sur-
rounded by setae, with hyaline, septate periphyses. Setae
1.5–3.5µm wide, numerous, cylindrical, erect, septate,
apex rounded, brown and tapering lighter at the apex.
Peridium 18–35µm (x̄ = 27µm, n = 20) wide, two layers,
outer layer composed of brown cells of textura angularis;
inner layer composed of hyaline cells of textura prismat-
ica. Asci 160–225 × 25–55 µm (x̄ = 190 × 40µm, n = 30),
8-spored, unitunicate, subglobose to clavate, pedicellate,
apex rounded, with J- apical ring, evanescent. Ascospores
27–35 × 15–25 µm (x̄ = 30 × 20 µm, n = 50), bi-seriate,
ellipsoidal to subglobose, brown to dark brown, aseptate,
with plate-like wrinkled wall and an apical germ pore, gut-
tulate. Asexual morph: Undetermined.
Material examined: Venezuela, Edo. Scure, NW of Irapa,
trail between Manacal and Los Pocitos, on dung of burro, 13
July 1972, K.P. Dumont, R.F. Cain, G.J. Samuels, G. Morillo
and J. Farfan (TRTC-175786, holotype).
Known hosts and distribution: On dung of burro in Ven-
ezuela (type locality) (Jeng and Krug 1976).
Notes: Emblemospora monotrema is similar to Neuros-
pora species and is characterized by pyriform ascomata and
clavate asci with one-celled ornamented ascospores (Jeng
and Krug 1976).
Eosphaeria Höhn., Annls mycol. 15(5): 362 (1917)
Isolated from soil. Sexual morph: Ascomata perithecial,
gregarious, superficial, globose to subglobose, brown to black,
carbonaceous, verrucose, ostiolate, papillate, periphysate.
Peridium carbonaceous to membranaceous, composed of
black to pale brown to hyaline cells of textura angularis to
textura prismatica. Paraphyses numerous, filiform, evanes-
cent. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex
rounded, with subapical globule, evanescent. Ascospores bi-
seriate, slightly curved, aseptate, multi-guttulate, hyaline and
Fig. 56 Diffractella curvata: (G-352276/1, holotype). a Herbarium
material label. b Herbarium material. c Ascoma on host. d Ascoma
surrounded by long setae. e Crushed ascoma. f Septate seta. g Asci.
h–k Ascospores stained in KOH. Scale bars: d = 500µm, e = 100µm,
f = 50µm, g–k = 20µm
Fungal Diversity
1 3
ellipsoidal when young, becoming multi-septate, geniculate,
2/3 upper cell swelling, truncate at the base, dark brown when
mature; 1/3 lower cell hyaline when mature, smooth-walled,
with bipolar short appendages, disappearing with age. Asexual
morph: Hyphomycetous, phialophora-like. Conidia small,
oval to globose (adapted from von Höhnel 1917).
Notes: The monotypic genus Eosphaeria is typified by E.
uliginosa and is characterized by subglobose ascomata and
L-shaped ascospores with short appendage at each end (von
Höhnel 1917). The genus was considered similar to Lasio-
sphaeria, and was thus initially accommodated in Lasio-
sphaeriaceae (Barr 1990; Kirk etal. 2001; Huhndorf etal.
2004b). Except for appendages at each end in ascospores,
Eosphaeria is similar to Immersiella in having black asco-
mata, cylindrical asci with an apical globule, and L-shaped
ascospores. Molecular data is unavailable for Eosphaeria.
Type species: Eosphaeria uliginosa (Fr.) Höhn., Annls
mycol. 15(5): 362 (1917)
Basionym: Sphaeria uliginosa Fr., in Kunze & Schmidt,
Mykologische Hefte (Leipzig) 2: 39 (1823)
Facesoffungi number: FoF 10041; Fig.58
Isolated from soil. Sexual morph: Ascomata
280–620µm (x̄ = 480µm, n = 10) diam., per ithecial, gre-
garious, superficial, globose to subglobose, brown to
black, carbonaceous, verrucose, ostiolate, with papilla,
periphysate. Peridium 60–100µm (x̄ = 75 µm, n = 20)
wide, comprising two layers, outer layer carbonaceous,
composed of black cells; inner layer membranaceous,
composed of pale brown to hyaline cells of textura angu-
laris to textura prismatica. Paraphyses numerous, fili-
form, evanescent. Asci (170–)190–250(–280) × 13–17µm
(x̄ = 225 × 16 µm, n = 20), 8-spored, unitunicate, cylin-
drical, pedicellate, apex rounded, with subapical glob-
ule, evanescent. Ascospores bi-seriate, slightly curved,
aseptate, multi-guttulate, hyaline and ellipsoidal when
young, becoming multi-septate, geniculate, 2/3 upper
cells swelling, truncate at the base, dark brown when
mature, 30–35(–45) × 4–7 µm (x̄ = 34 × 5.5µm, n = 50);
1/3 lower cells hyaline when mature, 12–18(–20) × 4–6µm
(x̄ = 15 × 4.5µm, n = 50), smooth-walled, with bipolar shor t
conical appendages (upper cauda 3.5–5.5 × 1–2.5µm; lower
Fig. 57 Emblemospora
monotrema: a, b (redrawn
from Jeng and Krug 1976); c–n
(TRTC-175786, holotype). a,
i Asci. b, k–n Ascospores. c
Herbarium material. d Ascoma
on host. e Squashed ascoma
f Ascoma in cross section. g
Peridium. h Setae at the neck. j
Apical structure in ascus. Scale
bars: a. h = 50µm, b = 20µm,
d = 500µm, e = 200µm, f.
i = 100µm, k–n = 10µm
Fungal Diversity
1 3
cauda 6.5–10 × 1.5–3µm), disappearing with age. Asexual
morph: Hyphomycetous, phialophora-like. Conidia small,
oval to globose (adapted from von Höhnel 1917).
Material examined: USA, Maine, Hancock Co. Township
of 728MD, Lead Mountain (Humpback), along road near
parking area, 44° 51ʹ 28″ N, 68° 05′ 52″ W, beech-maple for-
est, in soil, 2 September 2014, R.C. Harris (NY-01818643).
Known hosts and distribution: On soil in marshy places
in Germany (type locality) (Fries 1823b; von Höhnel 1917);
on clay soil in USA (Ellis and Everhart 1893).
Notes: Lasiosphaeria dichroospora, which was collected
on clay soil in USA, was synonymized under Eosphaeria
uliginosa (Ellis and Everhart 1893; von Höhnel 1917). We
could not obtain type material of E. uliginosa and, therefore,
re-examined an authentic specimen collected from USA by
Harris with features similar to those described in Ellis and
Everhart (1893) and von Höhnel (1917).
Isia D. Hawksw. & Manohar., Trans. Br. mycol. Soc. 71(2):
332 (1978)
Fig. 58 Eosphaeria uliginosa: a–f, h–m (NY-01818643); g (redrawn
from von Höhnel 1917). a Herbarium material label. b Herbarium
material. c Ascomata on host. d Ascoma in cross section. e Perid-
ium. f Asci with paraphyses. g Ascus. h–m Ascospores. Scale bars:
c = 1mm, d = 200µm, e–f = 100µm, g = 50µm, h–m = 20µm
Fungal Diversity
1 3
Saprobic on leaves or isolated from soil. Sexual morph:
Ascomata perithecial, solitary, immersed, membranaceous,
globose, dark brown to black, ostiolate, with papilla. Perid-
ium composed of membranaceous, brown to black cells of
textura angularis. Asci 8-spored, unitunicate, cylindrical,
evanescent. Ascospores uni-seriate, subglobose to ellipsoid,
0–1-septate, hyaline to pale brown, with dense irregular ver-
rucose ornamentation on surface. Asexual morph: Unde-
termined (adapted from Hawksworth and Manoharachary
1978; Udagawa and Sugiy 1982).
Notes: Hawksworth and Manoharachary (1978) intro-
duced Isia with I. neocaledoniensis as the type. It was trans-
ferred from Thielavia, and is similar to Copromyces and Jug-
ulospora in having ascospores with verrucose ornamentation
on the surface, although Copromyces has globose ascospores
and Jugulospora has a conical pedicel on ascospores (Hawk-
sworth and Manoharachary 1978). Therefore, this genus is
accommodated in Sordariales genera incertae sedis (Hawks-
worth and Manoharachary 1978; Maharachchikumbura etal.
2016; Hyde etal. 2020).
Type species: Isia neocaledoniensis (C. Moreau) D.
Hawksw. & Manohar., Trans. Br. mycol. Soc. 71(2): 334
(1978)
Basionym: Thielavia neocaledoniensis C. Moreau, Bull.
trimest. Soc. mycol. Fr. 81(4): 699 (1966)
Facesoffungi number: FoF 10042; Fig.59
Saprobic on leaves. Sexual morph: Ascomata peri-
thecial, solitary, immersed, membranaceous, globose,
dark brown to black, ostiolate, with papilla, periphysate.
Peridium composed of membranaceous, brown to black
cells of textura angularis. Asci (118–)120–135(–150) ×
13–18(–22) µm (x̄ = 130 × 16µm, n = 20) ascospore-bear-
ing part, 8-spored, unitunicate, cylindrical, apex rounded
or truncate, with J- apical ring. evanescent. Ascospores
18–23(–25) × (7.5–)9–12(–14) µm (x̄ = 20 × 10µm, n = 50),
uni-seriate, subglobose to ellipsoid, 0–1-septate, hyaline
when young, becoming pale brown at maturity, apex cuspi-
dal and base blunt, wall with densely verrucose ornamenta-
tion. Asexual morph: Undetermined.
Material examined: India, Bihar, on leaves of Pandanus
sp., 23 April 1977, M.A. Rizwi (IMI-213076, holotype).
Known hosts and distribution: On leaves of Pandanus in
New Caledonia and India (type locality) (Hawksworth and
Manoharachary 1978).
Notes: Thielavia neocaledoniensis was transferred to Isia
which was considered as a member of Sordariaceae (Hawk-
sworth and Manoharachary 1978). We re-examined the type
(213,076) that contains a permanent mount but we could find
only a few ascospores in its ascomata.
Lockerbia K.D. Hyde, Sydowia 46(1): 23 (1994)
Saprobic on palm fronds or wood. Sexual morph: Asco-
mata cleistothecial, solitary, superficial, membranaceous,
globose, dark brown to black. Peridium composed of brown
to hyaline cells of textura angularis to textura prismatica.
Paraphyses numerous, filamentous, branched, embedded in
a gelatinous matrix. Asci 8-spored, unitunicate, cylindrical
to subclavate, pedicellate, apex rounded and thick-walled.
Ascospores uni-seriate, limoniform to oval, hyaline to
brown to dark brown when mature, wall minutely verru-
cose, aseptate, guttulate, surrounded by inconspicuous hya-
line mucilaginous sheath. Asexual morph: Undetermined
(adapted from Hyde 1994; Raja and Shearer 2008).
Notes: Lockerbia palmicola was introduced as generic
type, and is similar to Diplogelasinospora, Neurospora
and Sordaria species in having ellipsoidal ascospores with
slightly irregular verrucose wall and surrounded by a hyaline
mucilaginous sheath (Hyde 1994; Raja and Shearer 2008).
Hence, Lockerbia was considered as a member in Sordari-
ales genera incertae sedis (Maharachchikumbura etal. 2016;
Hyde etal. 2020).
Type species: Lockerbia palmicola K.D. Hyde, Sydowia
46(1): 24 (1994)
Facesoffungi number: FoF 10043; Fig.60
Saprobic on palm fronds. Sexual morph: Ascomata
220–390µm (x̄ = 280µm, n = 5) diam., cleistothecial, soli-
tary, superficial, membranaceous, globose, dark brown
to black. Peridium 15–23µm (x̄ = 20 µm, n = 20) wide,
comprising two layers, outer layer composed of brown to
black cells of textura angularis; inner layer composed of
hyaline cells of textura prismatica. Paraphyses 1–2µm
wide, numerous, filamentous, branched, embedded in a
gelatinous matrix. Asci (120–)145–160(–170) × (13–)16–2
3(–25) µm (x̄ = 150 × 20µm, n = 20), 8-spored, unitunicate,
cylindrical to subclavate, pedicellate, apex rounded and
thick-walled. Ascospores (16–)18–21(–25) × 10–16(–18)
µm (x̄ = 20 × 15µm, n = 50), uni-seriate, limoniform to oval,
hyaline when young, brown to dark brown when mature,
aseptate, guttulate, wall minutely verrucose, surrounded
by inconspicuous hyaline mucilaginous sheath. Asexual
morph: Undetermined.
Material examined: Australia, north Queensland,
Bamaga, ‘Lockerbie’ rainforest, on dead palm rachides
(Archontophoenix sp.) on forest floor, February 1992, K.D.
Hyde (BRIP-21334, holotype).
Known hosts and distribution: On dead rachides of palm
in Australia (type locality) (Hyde 1994).
Notes: Hyde (1994) found Lockerbia palmicola is char-
acterized by cleistothecial ascomata and ellipsoidal to oval
ascospores. We re-examined the type (BRIP-21334).
Fungal Diversity
1 3
Periamphispora J.C. Krug, Mycologia 81(3): 476 (1989)
Coprophilous. Sexual morph: Ascomata perithecial, soli-
tary to scattered, superficial to semi-immersed, globose to
subglobose, brown, membranaceous to coriaceous, tubercu-
late, surrounded by hairs, ostiolate, with papilla. Peridium
composed of pale brown to hyaline cells of textura por-
recta to textura angularis to textura prismatica. Paraphy-
ses numerous, filiform, septate, evanescent. Asci 8-spored,
unitunicate, cylindrical to clavate, pedicellate, apex rounded.
Ascospores bi-seriate, ellipsoidal to oval, aseptate, ends
rounded, with a germ pore at the apex, hyaline when young,
becoming brown to reddish brown, rough-walled, sur-
rounded by hyaline, gelatinous sheath. Asexual morph:
Undetermined (adapted from Krug 1989).
Notes: The monotypic genus Periamphispora is char-
acterized by globose to subglobose ascomata with hairs
and ellipsoidal ascospores irregularly ribbed on the sur-
face, which is similar to Fimetariella, Podospora and
Sordaria (Krug 1989). This genus was previously con-
sidered a member in Lasiosphaeriaceae (Krug 1989;
Fig. 59 Isia neocaledoniensis:
(IMI-213076, holotype). a
Herbarium material label. b
Immersed ascomata on host.
c Ascoma. d, e Peridium.
f–h Asci. i–k Ascospores.
Notes: d–h from permanent
mount. Scale bars: b = 1mm,
c = 100µm, d. f–h = 50µm,
e = 20µm, i–k = 10µm
Fungal Diversity
1 3
Maharachchikumbura etal. 2015, 2016; Wijayawardene
etal. 2020). This genus is also similar to Podospora and
Sordaria in having hairy ascomata, clavate to cylindri-
cal asci and ellipsoidal ascospores with sheath, however
molecular data are unavailable.
Type species: Periamphispora phacelodes J.C. Krug, Myco-
logia 81(3): 476 (1989)
Facesoffungi number: FoF 10044; Fig.61
Coprophilous. Sexual morph: Ascomata
650–700 × 360–380µm (x̄ = 670 × 370µm, n = 5), perithe-
cial, solitary to scattered, superficial to semi-immersed, glo-
bose to subglobose, brown, membranaceous to coriaceous,
tuberculate, ostiolate, with necks. Necks brown to dark
brown, surrounded by numerous, pale brown, filiform, sep-
tate, hairs 1.5–3.5µm wide. Peridium 33–55µm (x̄ = 43µm,
n = 30) wide, outer layer composed of pale brown to red-
dish brown cells of textura porrecta to textura angularis;
inner layer composed of hyaline cells of textura prismatica.
Paraphyses numerous, filiform, septate, evanescent. Asci
200–300 × 40–65µm (x̄ = 250 × 55µm, n = 30) ascospore-
bearing part, 8-spored, unitunicate, cylindrical to clavate,
pedicellate, about 100µm long, apex rounded. Ascospores
(30–)40–45(–50) × 20–25(–30) µm (x̄ = 42 × 23µm, n = 50),
bi-seriate, ellipsoidal to oval, aseptate, ends rounded, with
a germ pore at the apex, hyaline when young, becoming
brown to reddish brown, rough-walled, surrounded by hya-
line, gelatinous sheath, 2–5µm wide. Asexual morph:
Undetermined.
Material examined: Spain, Málaga, about 20kmW of
Málaga, Benalmádena (36°36’N, 4°34’W), on horse dung,
30 April 1971, P. Blaser (TRTC-55023, holotype).
Known hosts and distribution: On horse dung in Spain
(type locality) (Krug 1989).
Notes: We re-examined the type, Periamphispora phace-
lodes 55023, which was grown on dung extract agar medium
culture (Weitzman and Silva-Hutner 1967), and it is well
preserved.
Fig. 60 Lockerbia palmicola:
(BRIP-21334, holotype). a,
b Herbarium material label.
c Herbarium material. d, e
Ascomata. f Ascoma in cross
section. g Peridium. h Para-
physes and an immature ascus.
i Ascus. j Ascospores with
minute pitting. k–n Ascospores.
Notes: h–k stained in Congo red
reagent. Scale bars: d = 500µm,
e = 200µm, f = 100µm,
g. i = 50µm, h. j = 20µm,
k–n = 10µm
Fungal Diversity
1 3
Reconditella Matzer & Hafellner, Biblthca Lichenol. 37: 46
(1990)
Saprobic on lichen. Sexual morph: Ascomata perithe-
cial, solitary, superficial to semi-immersed, membrana-
ceous, pyriform to subglobose, dark brown to black, ver-
rucose, surrounded by brown, septate setae, ostiolate, with
papilla, periphysate. Peridium composed of brown to hyaline
cells of textura angularis to textura prismatica. Paraphyses
unbranched, filiform. Asci 4–8-spored, unitunicate, cylindrical,
pedicellate, apex rounded. Ascospores uni- or bi-seriate, ellip-
soidal to fusiform, slightly curved, 0–1-septate, hyaline to pale
brown, striate and/or verrucose, guttulate. Asexual morph:
Undetermined (adapted from Matzer and Hafellner 1990).
Fig. 61 Periamphispora
phacelodes: (TRTC-55023,
holotype). a Herbarium
material label. b Ascomata on
Weitzman and Silva-Hutner’s
agar medium. c Appearance
of ascoma with hyphae. d
Ascoma in cross section. e
Peridium. f Perithecial neck
with periphyses. g Hyphae. h
Asci. i–k Ascospores with germ
pore (arrowed and enlarged
the apical germ pore). Scale
bars: b = 1mm, c = 500µm,
d = 200µm, e–g = 50µm,
h = 100µm, i–k = 20µm
Fungal Diversity
1 3
Notes: The monotypic genus Reconditella is typified by R.
physconiarum which was initially considered as a member
of Sordariales (Matzer and Hafellner 1990). We re-examined
the type material and found that this genus is characterized
by ellipsoidal to broadly fusiform, 0–1-septate ascospores
with fine warts forming ridges, which is similar to Podos-
pora and Neurospora. Therefore, we place this genus in Sor-
dariales genera incertae sedis.
Type species: Reconditella physconiarum Hafellner &
Matzer, in Matzer & Hafellner, Biblthca Lichenol. 37: 47
(1990)
Facesoffungi number: FoF 10046; Fig.62
Saprobic on lichen. Sexual morph: Ascomata
270–350 × 200–275µm (x̄ = 320 × 255µm, n = 5), perithe-
cial, solitary, superficial to semi-immersed, membrana-
ceous, pyriform to subglobose, dark brown to black, verru-
cose, surrounded by brown, septate setae 3.5–6.5µm wide,
ostiolate, with papilla, periphysate. Peridium 40–60µm
(x̄ = 45µm, n = 20) wide, comprising two layers, outer layer
composed of brown to black cells of textura angularis;
inner layer composed of hyaline cells of textura prismat-
ica. Paraphyses 5–6µm wide, unbranched, filiform. Asci
(70–)95–110(–120) × 10–17(–20) µm (x̄ = 100 × 15 µm,
n = 20), 4–8-spored, unitunicate, cylindrical, pedicellate,
apex rounded. Ascospores 15–20(–25) × 6–8(–9.5) µm
(x̄ = 18 × 7.5µm, n = 50), uni- or bi-seriate, ellipsoidal to
fusiform, slightly curved, 0–1-septate, hyaline to pale
brown, striate and/or verrucose, guttulate. Asexual morph:
Undetermined.
Material examined: Austria, Steiermark, Gesäuse area,
Johnsbach, c. 0.5km E of Gasthof Kölbl. Alt. c. 875m.
MTB 8453/4, on Fraxinus excelsior, on thallus of Physco-
nia distorta, 20 May 1988, S.J. Hafellner and E. Schreiner
(GZU-287606, holotype); ibid. (S-F20696, isotype).
Known hosts and distribution: On Physconia distorta in
Austria (type locality) (Matzer and Hafellner 1990).
Notes: We re-examined the type specimens of Recon-
ditella physconiarum. Specimen 287606 has fragile asco-
mata and has good quality permanent mount; F20696 is
well preserved. There is no molecular data available for R.
physconiarum.
Roselliniopsis Matzer & Hafellner, Biblthca Lichenol. 37:
97 (1990)
Saprobic on wood or lichens. Sexual morph: Ascomata
perithecial, solitary, superficial to semi-immersed, mem-
branaceous, pyriform to subglobose, black, verrucose,
ostiolate, with papilla. Peridium composed of pale brown
to hyaline cells of textura angularis to textura prismatica.
Paraphyses evanescent. Asci 8-spored, unitunicate, cylin-
drical to ellipsoidal, pedicellate, apex rounded. Ascospores
uni-seriate, ellipsoidal to fusiform, aseptate, pale brown to
dark brown, smooth-walled, with guttules. Asexual morph:
Undetermined (adapted from Matzer and Hafellner 1990;
Matzer 1993; Alstrup etal. 1994; Kondratyuk etal. 2013).
Notes: Roselliniopsis was established as a lichenicol-
ous genus in Sordariales and is typified by R. groedensis
(Matzer and Hafellner 1990; Maharachchikumbura etal.
2016; Hyde etal. 2020; Wijayawardene etal. 2020). It is
placed in Sordariales genera incertae sedis and characterized
by superficial ascomata with brown setae and pale brown
ascospores (Matzer and Hafellner 1990), but no molecular
data is available.
Type species: Roselliniopsis groedensis (Zopf) Matzer &
Hafellner, Biblthca Lichenol. 37: 99 (1990)
Basionym: Rosellinia groedensis Zopf, Hedwigia 35(6):
350 (1896)
Facesoffungi number: FoF 10047; Fig.63
Lichenicolous. Sexual morph: Ascomata
300–370 × 300–400µm (x̄ = 330 × 350µm, n = 5), per ithe-
cial, solitary, superficial to semi-immersed, membrana-
ceous, pyriform to subglobose, black, verrucose, ostio-
late, with papilla. Setae 5–10µm wide, brown, septate,
apex rounded, cells swollen near the septum. Peridium
50–120µm (x̄ = 75µm, n = 20) wide, comprising two lay-
ers, outer layer composed of pale brown cells of textura
angularis; inner layer composed of hyaline cells of textura
prismatica. Paraphyses evanescent. Asci (100–)110–12
5(–132) × (10–)14–17(–22) µm (x̄ = 120 × 15µm, n = 20),
8-spored, unitunicate, cylindrical to ellipsoidal, pedicellate,
apex rounded. Ascospores 14–20(–22) × (8–)10–12(–14)
µm (x̄ = 18 × 11 µm, n = 50), uni-seriate, oval to fusi-
form, aseptate, pale brown to dark brown, smooth-walled,
with germ pore at each end, guttulate. Asexual morph:
Undetermined.
Material examined: Italy, Trentino-Alto Adige, Bolzano,
St. Ulrich in Gröden (Südtirol), on Pertusaria sulphurella
Köerb. var. variolosa, W. Zopf (S-F47825).
Known hosts and distribution: On lichen in Italy (type
locality) (Zopf 1896; Matzer and Hafellner 1990).
Notes: Zopf (1896) introduced lichenicolous Rosellinia
groedensis and Matzer and Hafellner (1990) re-examined
its isotype from Italy and designated it as the type species
of Roselliniopsis. We re-examined an authentic specimen
collected by Zopf and found it to be similar to Podospora
and Sordaria in having membranaceous, black ascomata and
cylindrical asci with smooth-walled, ellipsoidal to fusiform
ascospores, but it lacks molecular data.
Synaptospora Cain, Beih. Sydowia 1: 4 (1957) [1956]
Saprobic on wood. Sexual morph: Ascomata perithecial,
superficial, gregarious to scattered, globose to subglobose,
carbonaceous, with ostiole, black, verrucose, surrounded by
setae. Peridium carbonaceous to membranaceous, composed
Fungal Diversity
1 3
Fig. 62 Reconditella physconiarum: a, b, e–g, i–n (S-F20696, iso-
type); c, d, h (GZU-287606, holotype). a Host lichen of Physconia
distorta. b Ascoma on host. c, e Ascomata in cross section. d, f, g
Peridium. h–j Asci (i with paraphyses). k–n Ascospores. Notes: c. d
from permanent mount. Scale bars: c. e = 100µm, d. f. g. i. j = 50µm,
h = 20µm, k–n = 10µm
Fungal Diversity
1 3
of hyaline to brown cells of textura angularis. Paraphyses
septate, cylindrical, evanescent. Asci 2–8-spored, unituni-
cate, cylindrical to clavate, apex broadly rounded or truncate.
Ascospores uni- or bi-seriate, globose to subglobose, aseptate,
hyaline when young, becoming pale brown to dark brown,
fused in groups. Asexual morph: Undetermined (adapted
from Cain 1957b; Huhndorf etal. 1999b; Réblová 2002).
Notes: Cain (1957b) established Synaptospora and
introduced two distinctive taxa, S. petrakii (type) and
S. tartaricola. The genus has mature ascospores which
become fused in groups and Cain (1957b) inferred that
this genus is closely related to Bombardia, Podospora and
Sordaria, due to its carbonaceous ascomata. Subsequently,
S. olandica, S. plumbea and S. setosa were introduced,
all of which have fused ascospores, except for S. plum-
bea with unfused, ellipsoidal ascospores (Huhndorf etal.
1999b; Réblová 2002). Miller etal. (2014) sequenced
S. plumbea (ANM963 and SMH3962) and found that it
Fig. 63 Roselliniopsis groeden-
sis: (S-F47825). a Herbarium
material label. b Herbarium
material. c Ascomata on host.
d Ascoma in cross section. e
Peridium. f Part of the ostiole
with periphyses. g Seta. h–j
Asci. k–n Ascospores. Scale
bars: d = 100µm, e–j = 50µm,
k–n = 10µm
Fungal Diversity
1 3
clustered with Helminthosphaeria species. Thus, Synapto-
spora was transferred to Helminthosphaeriaceae based
on phylogenetic analyses (Miller etal. 2014; Hyde etal.
2020; Wijayawardene etal. 2020). We found that S. plum-
bea is similar to Helminthosphaeria in having globose
ascomata, cylindrical asci and brown aseptate, ellipsoidal,
unfused ascospores, differing from other Synaptospora
species with fused ascospores. Phylogenetically, S. plum-
bea nested in Helminthosphaeriaceae (61%ML, Fig.21).
Therefore, we recommend transferring S. plumbea to Hel-
minthosphaeria. The remaining four Synaptospora species
have globose ascomata, cylindrical asci and globose, fused
ascospores, but no molecular information is available. We
place Synaptospora in Sordariales genera incertae sedis
pending future work.
Type species: Synaptospora petrakii Cain, Beih. Sydowia
1: 5 (1957) [1956]
Facesoffungi number: FoF 10048; Fig.64
Saprobic on decorticated wood. Sexual morph: Ascomata
200–350 × 250–450µm (x̄ = 300 × 350µm, n = 5), perithecial,
superficial, gregarious to scattered, globose to subglobose,
carbonaceous, with ostiole, black, verrucose, surrounded by
sparse, brown, septate setae, 3.5–7µm wide, with a rounded
apex. Subiculum ascomata seated on sparse, brown, septate,
branched, friable hyphae, 5.5–7.5µm wide. Peridium 30–75µm
(x̄ = 55µm, n = 20) wide, comprising two layers, outer layer car-
bonaceous, composed of brown cells of textura angularis; inner
layer membranaceous, composed of hyaline cells of textura
prismatica. Paraphyses 2.5–8.5µm wide, septate, slender to
cylindrical, evanescent. Asci (70–)80–95(–100) × 7.5–10µm
(x̄ = 85 × 8µm, n = 30), normally 2-spored, unitunicate, cylindri-
cal, broadly rounded and thickened at the apex, long pedicellate.
Ascospores (20–)22–24(–26) × 4–8µm (x̄ = 23 × 6µm, n = 50)
for four cells, uni-seriate, subglobose to ellipsoidal, hyaline
and aseptate when young, gradually becoming cylindrical to
oblong, dark brown, usually 2–4-septate, distinctly constricted
at the septa, the cells swollen, ends broadly rounded, straight
to slightly curved, smooth-walled, with a large guttule in each
cell, breaking into part spores. Part spores 6–9 × 4.5–7 µm
(x̄ = 7.5 × 5.5µm, n = 50), hyaline to dark brown, globose to
ellipsoidal, aseptate, smooth-walled, with a large guttule, dis-
charging through constricting septa. Asexual morph: Undeter-
mined (adapted from Cain 1957b).
Material examined: Canada, Ontario, north of Missis-
sauga, in willow grove by Credit River on Creditview Road,
on decaying logs, 11 November 1987, L.A. Novak (TRTC-
51203); Canada, Ontario, Muskoka district, Haliburton,
11km south of Dorset, on decorticated wood, October 1975,
D. Tighe (TRTC-51205).
Known hosts and distribution: On dead decorticated log
of Betula papyrifera in Canada (type locality) (Cain 1957b).
Notes: The holotype material (TRTC-32168) was col-
lected by Cain (1957b) in Canada. We were unable to
obtain the type material and, therefore, we re-examined two
authentic samples, TRTC-51203 and TRTC-51205, col-
lected in Canada and determined by M. Matzer (mentioned
in the label of material). Ascospores of Synaptospora was
described as ellipsoidal to globose and fused in groups when
mature (Cain 1957b; Barr 1990; Huhndorf etal. 1999b).
We could not find mature ascospores fused in groups in
TRTC-51203 and TRTC-51205. However, we found that
the ascospores were initially hyaline, ellipsoidal to oblong
or cylindrical and gradually formed distinct septa, and finally
the septa split the brown ascospores into individual globose
to subglobose part spores (Fig.64m–r). Fresh collections of
Synaptospora species are required to determine the nature
of ascospores and placement of the genus.
Tripterosporella Subram. & Lodha, Curr. Sci. 37: 246
(1968)
Coprophilous Sexual morph: Ascomata cleistothe-
cial, scattered, superficial, globose to subglobose, semi-
translucent to opaque, dark brown to black, surrounded
by brown, septate hairs. Peridium membranaceous, com-
posed of brown cells of textura angularis to textura pris-
matica. Asci 8-spored, unitunicate, clavate to cylindrical,
pedicellate, evanescent, apex rounded or truncate, with
J-apical ring. Paraphyses filiform or absent. Ascospores
bi-seriate or irregularly arranged, cylindrical, aseptate
and hyaline when young, upper cell(s) becoming swollen
and fusiform, 0–1-septate, brown to olivaceous brown,
guttulate, with truncate base and an apical germ pore,
collapsing when dry; lower cell(s) tail-like, hyaline,
aseptate to multi-septate, smooth-walled, slightly curved
near the base. Asexual morph: Undetermined (adapted
from Subramanian and Lodha 1968; Abdullah and Rat-
tan 1978).
Notes: Tripterosporella was isolated from herbivores
dung (horses and cows), and is similar to lasiosphaeriaceous
taxa in having ascospores with brown upper cell(s) and a
long hyaline pedicel (Subramanian and Lodha 1968; Doveri
2010; Wijayawardene etal. 2020). We place Tripterosporella
in Sordariales genera incertae sedis but phylogenetic data is
required to confirm its placement.
Type species: Tripterosporella coprophila Subram. &
Lodha, Curr. Sci. 37: 246 (1968)
Facesoffungi number: FoF 10138; Fig.37f–g
Coprophilous Sexual morph: Ascomata cleistothecial,
scattered, superficial, globose, black, membranaceous, sur-
rounded by brown, septate hairs. Peridium membranaceous,
Fungal Diversity
1 3
composed of brown to hyaline cells of textura angularis to
textura prismatica. Asci 8-spored, unitunicate, clavate to
cylindrical, pedicellate, apex rounded, with J- apical ring.
Ascospores bi-seriate, cylindrical, aseptate and hyaline
when young, upper cell(s) becoming swollen and ellipsoi-
dal to subglobose, 0–1-septate, brown, with truncate base
and an apical germ pore, collapsing when dry; lower cell(s)
tail-like, hyaline, 0–mulit-septate, smooth-walled, slightly
curved near the base. Asexual morph: Undetermined
(adapted from Abdullah and Rattan 1978).
Known hosts and distribution: On dung of Bos taurus in
India (type locality) and Iraq (Subramanian and Lodha 1968;
Abdullah and Rattan 1978).
Notes: Tripterosporella coprophila has brown ascospores
with ellipsoidal upper cell(s) and a long, cylindrical, hya-
line, multi-septate pedicel (Subramanian and Lodha 1968;
Abdullah and Rattan 1978). However, the septate pedicel
was rarely mentioned in subsequent researches (Abdullah
and Rattan 1978; Doveri 2010). Tripterosporella coproph-
ila is similar to Podospora, Lasiosphaeria and Zopfiella in
Fig. 64 Synaptospora petrakii:
a–c, e, f, h–o, q–x (TRTC-
51203); d, g, p (TRTC-51205).
a Herbarium material. b
Ascomata on host. c, d Ascoma
in cross section. e Peridium. f, g
Setae. h Hyphae of subiculum.
i Septate paraphyses. j Asci
with paraphyses. k Asci. l Apex
of ascus (stained in Melzer's
reagent). m–q, s–v Immature
to mature ascospores (m–q
3-septate; s–t 2-septate; u–v
4-septate). r, w Ascospores
breaking into individual cells.
x Individual spore cells.
Scale bars: b = 500µm, c.
d. j = 100µm, e. k = 50µm,
f–i = 20µm, l–x = 10µm
Fungal Diversity
1 3
having cleistothecial ascomata and multi-septate ascospores
comprising two types of cells. We were unable to obtain
the authentic material of this species and therefore, we
refer to Subramanian and Lodha (1968) for hand-drawing
(Fig.37f–g).
Other genera studied
Subclass Xylariomycetidae O.E. Erikss. & Winka
Xylariales Nannf.
Diatrypaceae Nitschke
Monosporascus Pollack & Uecker, Mycologia 66(2): 348
(1974)
Saprobic on wood. Sexual morph: Colonies white and
hyaline. Ascomata perithecial, solitary to gregarious, semi-
immersed, membranaceous, ellipsoidal to globose, dark
brown to black, ostiolate, with papilla. Peridium mem-
branaceous, comprising brown to olivaceous-brown cells
of textura angularis to textura epidermoidea. Paraphyses
numerous, filiform, cylindrical, flexuous, septate. Asci 1, 2,
3 or 4-spored, unitunicate, clavate to cylindrical, pedicellate.
Ascospores globose, aseptate, hyaline to bright brown to
brown to black, smooth-walled, guttulate. Asexual morph:
Undetermined (adapted from Pollack and Uecker 1974;
Negreiros etal. 2019).
Notes: Pollack and Uecker (1974) introduced Monos-
porascus typified by M. cannonballus. It has a Xylaria-
type centrum and 1 or 2-spored asci with aseptate, glo-
bose ascospores with guttules. This genus was shown to
belong to Diatrypaceae (Xylariales) based on phyloge-
netic analyses (Schoch etal. 2014; Maharachchikumbura
etal. 2016; Negreiros etal. 2019; Hyde etal. 2020; this
study, Fig.1).
Type species: Monosporascus cannonballus Pollack &
Uecker, Mycologia 66(2): 348 (1974)
Facesoffungi number: FoF 10049; Fig.65
Culture on malt yeast agar from Cucumis melo. Sex-
ual morph: Colonies white and hyaline. Ascomata
310–500 × 270–420µm (x̄ = 380 × 330µm, n = 10), per i-
thecial, solitary to gregarious, semi-immersed, membrana-
ceous, ellipsoidal to globose, dark brown to black, ostiolate,
with papilla. Peridium 35–80µm (x̄ = 55µm, n = 20) wide,
comprising 5-layers, outer layer composed of hyaline, plect-
enchymatous tissues; next layer composed of olive-brown
cells of textura epidermoidea; third layer composed of hya-
line cells of textura angularis cells; fourth layer composed
of olive-brown to brown cells of textura angularis; inner
layer composed of hyaline cells of textura prismatica. Para-
physes 3–6µm wide, numerous, filiform, cylindrical, flexu-
ous, septate. Asci (40–)45–53(–60) × (18–)20–30(–35) µm
(x̄ = 50 × 24µm, n = 20), 1-spored, unitunicate, clavate to
pyriform, thick-walled, pedicellate, persistent. Ascospores
1-celled, globose, hyaline when young, becoming bright
brown to brown (15–25µm diam.) to black (35–45µm
diam.) at maturity, smooth-walled, guttulate. Asexual
morph: Undetermined.
Material examined: USA, Arizona, Yuma, from Cucumis
melo L., on culture of malt yeast agar-1/4 dilution, 21 June
1972, F.G. Pollack and F.A. Uecker (BPI-579840, holotype).
Known hosts and distribution: On Cucumis melo in Bra-
zil, Spain and USA (type locality) (Pollack and Uecker 1974;
Schoch etal. 2014).
Notes: We re-examined the type of Monosporascus can-
nonballus (BPI-579840). The strain of M. cannonballus is
sister to M. mossoroensis and M. nordestinus (Diatrypaceae)
(Fig.1).
Sordariomycetes genera incertae sedis
Conidiotheca Réblová & L. Mostert, Mycol. Res. 111(3):
305 (2007)
Saprobic on wood. Sexual morph: Ascomata perithe-
cial, solitary to scattered, immersed to semi-immersed,
dark brown to black, subglobose to broadly conical, ostio-
late, with papilla, periphysate. Peridium membranaceous,
comprising two layers, outer layer composed of black to
pale brown cells of textura angularis; inner layer composed
of hyaline cells of textura prismatica. Paraphyses numer-
ous, septate, tapering, longer than the asci. Asci 8-spored,
cylindrical to clavate, short pedicellate, apex obtuse, with
J- apical ring. Ascospores 1–2-seriate, hyaline, ellipsoidal
to subglobose, 0–1-septate, sometimes slightly curved when
young, becoming fusiform, with transverse and longitudi-
nal septa, constricted at septa, producing several lateral
appendices as ascoconidia. Ascoconidia hyaline, allantoid
to ellipsoidal, aseptate, smooth-walled, guttulate, filling the
whole ascus and discharged through constricted apical ring.
Asexual morph: Undetermined (adapted from Réblová and
Mostert 2007).
Notes: Barr (1985) described Romellia tympanoides (≡
Conidiotheca tympanoides) with clavate asci and ascospores
producing ascoconidia similar to Tympanis (Leotiomyceti-
dae). Barr etal. (1993) transferred it to Jattaea based on the
spicate clusters of clavate asci. Réblová and Mostert (2007)
transferred J. tympanoides to Conidiotheca which was placed
in Ascomycota genera incertae sedis. Maharachchikumbura
etal. (2015, 2016) and Wijayawardene etal. (2018) placed
this genus in Calosphaeriales and Togniniales, respectively.
We re-examined the type and found that Conidiotheca is sim-
ilar to Jattaea (Calosphaeriaceae, Diaporthomycetidae) and
Pleonectria (Nectriaceae, Hypocreomycetidae), which have
clavate asci and septate ascospores with numerous ascoco-
nidia. However, Conidiotheca has brown to black ascomata
whereas those of Pleonectria are yellow to red (Réblová and
Fungal Diversity
1 3
Mostert 2007; Hirooka etal. 2012). Conidiotheca tympa-
noides is also similar to Jattaea taediosa, as they both have
black ascomata, clavate asci and septate ascospores with
numerous ascoconidia when mature, however, the former
does not have the ascogenous hyphae unique to the latter
(Réblová etal. 2015). Unfortunately, there is no molecular
data available for C. tympanoides. We place this genus in
Sordariomycetes genera incertae sedis pending more data.
Type species: Conidiotheca tympanoides (M.E. Barr)
Réblová & L. Mostert, Mycol. Res. 111(3): 305 (2007)
Fig. 65 Monosporascus cannonballus: (BPI-579840, holotype). a Dry culture. b Ascoma with collapsing papilla. c, d Vertical sections of asco-
mata. e Peridium. f Paraphyses. g–j Asci. k–n Ascospores. Scale bars: b–d = 200µm, e = 50µm, f–n = 20µm
Fungal Diversity
1 3
Basionym: Romellia tympanoides M.E. Barr, Mycologia
77(4): 561 (1985)
Facesoffungi number: FoF 10050; Fig.66
Saprobic on wood. Sexual morph: Ascomata
250–455 × 350–550µm (x̄ = 345 × 425µm, n = 10), per i-
thecial, solitary to scattered, papillate, immersed, erumpent
through the bark of the host, dark brown to black, subglo-
bose to broadly conical, ostiolate, with papilla, periphysate.
Peridium 25–50µm (x̄ = 35µm, n = 30) wide, comprising
two layers, outer layer composed of black to pale brown
cells of textura angularis; inner layer composed of hya-
line cells of textura prismatica. Paraphyses 2–3.5µm wide
(x̄ = 2.8µm, n = 30), numerous, septate, tapering, longer
than the asci. Asci (120–)150–180(–200) × 15–18(–25) µm
(x̄ = 170 × 16µm, n = 50), 8-spored, cylindrical to clavate,
short pedicellate, apex obtuse, with J-apical ring. Ascospores
(10–)13–20(–25) × 3.5–6.5 µm (x̄ = 18 × 5 µm, n = 50),
1–2-seriate, hyaline, ellipsoidal to subglobose, 0–1-septate,
sometimes slightly curved when young, becoming fusiform,
with transverse and longitudinal septa, constricted at septa,
producing several lateral appendices as ascoconidia. Asco-
conidia (2.5–)3–5(–6) × 1–2.5µm (x̄ = 4 × 1.5 µm, n = 50),
hyaline, allantoid to ellipsoidal, aseptate, smooth-walled,
guttulate, filling the whole ascus and discharged through
constricted apical ring. Asexual morph: Undetermined.
Material examined: USA, Massachusetts, Baptist Hill,
Conway, 21 April 1969, M.E. Barr, on twigs of Prunus
pennsylvanica (NY-00912076, holotype).
Known hosts and distribution: On decayed wood of Pru-
nus pennsylvanica in the USA (type locality) (Réblová and
Mostert 2007).
Notes: The monotypic Conidiotheca is typified by C.
tympanoides (Réblová and Mostert 2007). The holotype
NY00912076 is well preserved, but there are few complete
asci, and production of ascoconidia is unclear. Therefore,
the hand-drawing is provided for the asci (Fig.66l–n) and
the process of producing ascoconidia (Fig.66s–v) based on
Réblová and Mostert (2007).
Ascomycota genera incertae sedis
Copromyces N. Lundq., Ark. Bot. 6: 327 (1967)
Coprophilous. Sexual morph: Ascomata cleistothecial,
solitary, superficial, globose to subglobose, membranaceous,
dark brown, with hyaline surface hairs. Peridium membrana-
ceous, composed of dark brown to hyaline cells of textura
angularis. Asci 2-spored, ovoid, apex rounded, short pedi-
cellate. Ascospores globose to subglobose, bright yellow,
becoming olivaceous, brown to black at maturity, aseptate,
guttulate, with a germ pore and irregular plate-like wrin-
kles on the surface. Asexual morph: Undetermined (adapted
from Lundqvist 1967; Avila etal. 2002).
Notes: The monotypic genus Copromyces is typified by
C. bisporus and has cylindrical to clavate, 2-spored asci and
globose ascospores (Lundqvist 1967; Avila etal. 2002). The
genus was initially placed in Sordariaceae (Lundqvist 1967).
Petrak (1967) transferred C. bisporus to Rechingeriella (Zop-
fiaceae, Dothideomycetes), because it has 2-spored asci and
tubercular ascospores similar to those of R. insignis. Subse-
quently, Lundqvist (1972) suggested that Rechingeriella and
Copromyces are unrelated, because R. insignis has black, car-
bonaceous ascomata, distinctly bitunicate asci and ascospores
with no germ pore, whereas, C. bisporus has membranaceous
ascomata and ascospores with germ pore. Later, Avila etal.
(2002) recorded that C. bisporus has cleistothecial ascomata
and unitunicate asci with globose ascospores. Unfortunately,
molecular data for C. bisporus is unavailable.
Copromyces sp. (CBS 386.78) was reported in the taxa
sequence list of Huhndorf etal. (2004b) and noted as strain
TRTC 51747 (& CBS 386.78, named as C. octosporus Jeng
& Krug). This species, however, has not been formally
described and is invalid. ‘Copromyces octosporus’ (CBS
386.78) was listed as the second species in Copromyces, and
placed in Sordariaceae based on LSU and ITS sequence data
(Huhndorf etal. 2004b; Maharachchikumbura etal. 2015,
2016; Vu etal. 2019; Hyde etal. 2020). However, in this
study, the strain CBS 386.78 is sister to Pseudoneurospora
canariensis (91%ML, Fig.40). As the species has not been
formally described and there is no apparent holotype it needs
recollecting and introducing.
We re-examined the isotype (749020) of Copromyces
bisporus, which is well preserved, and observed that it has
inconspicuously evanescent asci and dissolved hyaline fila-
ments between the asci. We place Copromyces in Ascomy-
cota genera incertae sedis pending molecular data.
Type species: Copromyces bisporus N. Lundq., Arch. für
Botanik, Ser. 2 6: 328 (1967)
Facesoffungi number: FoF 10051; Fig.67
Coprophilous on rabbit dung. Sexual morph: Ascomata
200–250µm (x̄ = 235µm, n = 10) diam., cleistothecial, solitary,
uniloculate, superficial, globose to subglobose, membranaceous,
dark brown, with hyaline surface hairs. Peridium 30–50µm
wide, membranaceous, composed of dark brown to hyaline
cells of textura angularis. Asci (24–)26–32(–35) × 12–15(–17)
µm (x̄ = 30 × 13µm, n = 50), 2-spored, ovoid, apex rounded,
short pedicellate. Ascospores (11–)12–16(–20) µm (x̄ = 15µm,
n = 50) diam., globose to subglobose, bright yellow, becoming
olivaceous, brown to black at maturity, aseptate, guttulate, with
a germ pore and irregular plate-like wrinkles on the surface.
Asexual morph: Undetermined.
Material examined: Sweden, Skane, Ivetofta parish, Aby,
in pasture, on rabbit dung, 8 June 1962, N. Lundqvist (BPI-
749020, isotype).
Known hosts and distribution: On dung of rabbit (Orycto-
lagus cuniculus) in Sweden (type locality) (Lundqvist 1967);
on rabbit and fox dung in Venezuela (Avila etal. 2002).
Fungal Diversity
1 3
Effetia Bartoli, Maggi & Persiani, Mycotaxon 19: 517
(1984).
Isolated from forest soil. Sexual morph: Ascomata peri-
thecial, solitary, brown to dark brown, globose to subglo-
bose, ostiolate, with short necks. Peridium membranaceous.
Asci 8-spored, unitunicate, cylindrical, apex rounded.
Ascospores hyaline to brown, ellipsoidal, aseptate, with an
apical germ pore. Asexual morph: Hyphomycetous. Myce-
lium dark brown, superficial, forming a thick subiculum.
Hyphae flexuous, brown, septate, branched, becoming brit-
tle. Conidiophores brown, unbranched, straight or flexu-
ous, smooth-walled, septate, narrowed towards apex. Con-
idiogenous cells polyblastic, terminal ones give rise to new
ones becoming intercalary, sympodial, swollen at the apex.
Fig. 66 Conidiotheca tympanoides: a–k, o–r, w (NY-00912076, hol-
otype); l–n (redrawn from Réblová and Mostert 2007); s–v (Drawn
following observation by microscope). a Herbarium material. b Gre-
garious ascomata. c Ascomata. d Ascoma in cross section. e Perid-
ium. f, g Septate paraphyses. h Ascus with paraphyses. i, j Asco-
conidia filling the whole ascus and discharged through constricted
apical ring. k–n Ascospores gradually swollen in ascus and produc-
ing large numbers of ascoconidia. o–v Immature to mature ascospores
(p–r, t–v ascospores producing ascoconidia). w Ascoconidia.
Scale bars: c = 500 µm, d = 200 µm, e–j = 50 µm, k–n, w = 20 µm,
o–v = 10µm
Fungal Diversity
1 3
Conidia acropleurogenous, oval to ellipsoidal, aseptate,
light brown to dark brown, smooth-walled, with a sheath.
Chlamydospores single or like a string of beads, irregular,
brown, thick-walled (adapted from Bartoli etal. 1984).
Notes: Bartoli etal. (1984) introduced the monotypic
genus Effetia and proposed that its sexual morph is similar to
Apodus and Fimetariella based on its black ascomata, cylin-
drical asci and ellipsoidal, hyaline to light brown ascospores,
while its asexual morph (in culture) was described as vir-
gariella-like, which is more similar to Xylariaceae. However,
Bartoli etal. (1984) proposed to place Effetia in Sordari-
aceae based on its sexual morph, but no molecular data is
available. We recommend transferring Effetia to Ascomycota
genera incertae sedis pending future work.
Type species: Effetia craspedoconidica Bartoli, Maggi &
Persiani, Mycotaxon 19: 517 (1984)
Facesoffungi number: FoF 10052; Fig.68a–q
Isolated from forest soil. Sexual morph: Ascomata peri-
thecial, solitary, brown to dark brown, globose to subglobose,
ostiolate, with short necks. Peridium membranaceous. Asci
8-spored, unitunicate, cylindrical, apex rounded. Ascospores
hyaline to brown, ellipsoidal, aseptate, with a germ pore.
Asexual morph: Hyphomycetous. Mycelium dark brown,
superficial, forming a thick subiculum. Hyphae 1.5–3.5µm
(x̄ = 2.5µm, n = 30) wide, flexuous, brown, septate, branched,
becoming brittle. Conidiophores brown, unbranched, straight
or flexuous, smooth-walled, septate, narrowed towards apex.
Conidiogenous cells polyblastic, terminal ones give rise to
new ones becoming intercalary, sympodial, swollen at the
apex. Conidia 7.5–11 × 3–5µm (x̄ = 9.5 × 4µm, n = 50), acro-
pleurogenous, oval to ellipsoidal, aseptate, light brown to
dark brown, smooth-walled, with a sheath 1–1.5µm wide.
Fig. 67 Copromyces bisporus: (BPI-749020, isotype). a Herbarium
material label. b Ascoma. c Ascoma with hyaline hairs (arrow).
d Vertical section of ascoma. e Peridium. f Evanescent filaments
(stained in Melzer’s reagent). g–j Asci stained in 5% KOH (h. arrow
marks germ pore, h–j are the same ascus). Scale bars: b. c = 200µm,
d = 100µm, e = 50µm, f = 20µm, g–j = 10µm
Fungal Diversity
1 3
Chlamydospores (5–)7–10(–12) µm (x̄ = 9µm, n = 20) diam.,
single or like a string of beads, irregular, brown, thick-walled
(adapted from Bartoli etal. 1984).
Material examined: Ivory Coast, Tai forest National Park,
on rain forest litter, December 1976, B. Rambelli (IMI-
223521, holotype).
Known hosts and distribution: On woodland soil in Ivory
Coast (type locality) (Bartoli etal. 1984).
Notes: Bartoli etal. (1984) noted that the sexual morph
of Effetia craspedoconidica was produced only in the initial
isolates, and then it lost viability. We re-examined the type
Fig. 68 Effetia craspedoconidica: a–e (redrawn from Bartoli et al.
1984); f–q (IMI-223521, holotype). a Part of young ascomata on
surface view. b Vertical view of ascoma with asci. c Ascus. d Con-
idiophores with conidia. e, o–q Chlamydospores (o. young state, p–q.
dry and collapsed). f Part of colony. g, h Conidiophores with conidia
(h. arrow marks swollen and sympodial conidiogenous cells). i–n
Conidia with sheath; Boothiella tetraspora: r, s (redrawn from Lodhi
and Mirza 1962). r Ascus s Ascospores. Scale bars: b = 50µm, c. g.
h. o–s = 10µm, d. e = 20µm, f = 500µm, i–n = 5µm
Fungal Diversity
1 3
(223521) and only found the asexual morph in culture. We
draw the sexual morph with reference to Bartoli etal. (1984).
Endophragmiella B. Sutton, Mycol. Pap. 132: 58 (1973)
Sexual morph: Undetermined. Asexual morph: Hypho-
mycetous. Mycelium immersed to superficial, composed
of pale brown to brown, septate, branched, smooth-walled
hyphae. Conidiophores macronematous, mononematous,
erect, simple or branched, straight or flexuous, pale brown,
smooth-walled. Conidiogenous cells monoblastic, integrated,
terminal, tapering towards the truncate apex. Conidia soli-
tary, acrogenous, simple, ellipsoidal, pale brown to brown,
smooth-walled, 1–multi-septate (adapted from Sutton 1973).
Notes: The hyphomycete genus Endophragmiella is typi-
fied by End. pallescens which was found associated with
stromata of Cytospora chrysosperma (Valsaceae) on wood
of Populus tremuloides (Sutton 1973). Hughes (1979) sug-
gested that Endophragmiella maybe the sexual morph of
Echinosphaeria and the synanamorph of Selenosporella.
Hughes (1979) also described two specimens of Echino-
sphaeria canescens, one of which was accompanied by
End. biseptata; and the other was related to Selenosporella
(Hughes 1979). These associations are not confirmed
through culture or phylogenetic analysis. Molecular data is
available for End. dimorphospora (CBS 438.74) and End.
taxi (CBS 614.84), but not for End. pallescens (Hernández-
Restrepo etal. 2017; Vu etal. 2019). End. dimorphospora
nests in Helminthosphaeriaceae (Fig.21), but End. taxi is
closely related to members of Tubeufiales (Dothideomy-
cetes) (not shown).
Type species: Endophragmiella pallescens B. Sutton,
Mycol. Pap. 132: 62 (1973)
Facesoffungi number: FoF 10139; Fig.22s
Sexual morph: Undetermined. Asexual morph: Hypho-
mycetous. Colonies effuse, reddish brown, hairy, originating
around effete stromata of immersed microfungi and spread-
ing over the bark. Mycelium about 4µm wide, immersed to
superficial, composed of brown, septate, branched, smooth-
walled hyphae. Conidiophores macronematous, monon-
ematous, erect, simple or branched, flexuous, infrequently
rigid, septate, light brown, becoming pale brown towards the
apex, smooth-walled. Conidiogenous cells monoblastic, inte-
grated, cylindrical, terminal, apex with 1–3 unflared annel-
lations, pale brown. Conidia solitary, acrogenous, simple,
ellipsoidal, pale brown, smooth-walled, 1–2-septate, not
constricted at the septa, with a basal marginal frill (adapted
from Sutton 1973).
Known hosts and distribution: On dead wood of Populus
tremuloides in Canada (type locality) (Sutton 1973).
Notes: The mycelium of Endophragmiella pallescens
associated with stromata of Cytospora chrysosperma on
natural substrate in Manitoba, Canada (Sutton 1973). The
hand-drawing is provided for the type species (Fig.22s)
based on Sutton (1973).
Tulipispora Révay & Gönczöl, in Révay, Gönczöl & Des-
cals, Nova Hedwigia 88(1-2): 42 (2009)
Saprobic on wood, rarely on leaves. Sexual morph:
Undetermined. Asexual morph: Hyphomycetous. Myce-
lium semi-immersed in wood. Conidiophores hyaline, semi-
macronematous, mononematous, simple, septate. Conidiog-
enous cells integrated, terminal, proliferations sympodial.
Conidia solitary, acrogenous to subacrogenous, multi-sep-
tate, composed of elongate main axis and branches, branches
mostly subverticillate, sometimes grouped near the base of
the axis, curved (adapted from Révay etal. 2009).
Notes: The monotypic Tulipispora resembles Triscelo-
phorus and Triramulispora in having multi-septate conidia
with an elongate main axis and branches (Révay etal. 2009).
It has been accommodated in Calosphaeriaceae, Sordari-
omycetes and Ascomycota genera incertae sedis (Maharach-
chikumbura etal. 2015, 2016; Wijayawardene etal. 2018,
2020). However, as an asexual genus, Tulipispora is dis-
similar to the known asexual morph of other members in
Calosphaeriaceae. Tulipispora species have multi-septate
and elongate conidia, whereas species of Calospaheriaceae
usually have asepate, allantoid conidia. We place Tulipispora
in Ascomycota genera incertae sedis.
Type species: Tulipispora ingoldii Révay & Gönczöl,
Nova Hedwigia 88(1-2): 42 (2009)
Facesoffungi number: FoF 10140; Fig.69
Saprobic on wood, rarely on leaves. Sexual morph:
Undetermined. Asexual morph: Hyphomycetous. Myce-
lium semi-immersed in wood. Conidiophores hyaline,
semi-macronematous, mononematous, simple, septate.
Conidiogenous cells integrated, terminal, proliferations
sympodial. Conidia solitary, acrogenous to subacrog-
enous, multi-septate, composed of elongate main axis
and branches, branches mostly subverticillate, sometimes
grouped near the base of the axis, curved (adapted from
Révay etal. 2009).
Fungal Diversity
1 3
Known hosts and distribution: On decayed wood and
leaves in Hungary (type locality) (Révay etal. 2009).
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 Saranyaphat Boonmee, Putarak Chomnunti,
Dong-Qin Dai, Anusha H. Ekanayaka, Ji-Chuan Kang, Ming 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.
Declaration
Conflict of interest The authors declare that no known conflicts of in-
terests exist.
References
Abdullah SK (1983) New and noteworthy Ascomycetes from Iraq.
Trans Br Mycol Soc 81(2):392–396
Abdullah SK, Rattan SS (1978) Zygopleurage, Tripterosporella and
Podospora (Sordariaceae; Pyrenomycetes) in Iraq. Mycotaxon
7(1):102–116
Albertini JB, Schweinitz LD (1805) Conspectus Fungorum in Lusatiae
superioris, pp 1–376
Alstrup V, Christensen SN, Hansen ES etal (1994) The Lichens of the
Faroes. Fródskaparrit 40:61–121
Andersson K, Eriksson OE, Landvik S (1995) Boliniaceae transferred
to Sordariales (Ascomycota). Syst Ascomycetum 14:1–16
Arambarri AM, Minter TJ, Cabello MN etal (2020) Spegazzini's draw-
ings of fungi, a digitized library. www. cyber truffl e. org. uk/ spega zzini
Avila AED, Chávez AJP, García LMU (2002) Copromyces bisporus,
a new record from Venezuela. Revista Científica 12(1):12–14
Barr ME (1985) Notes on the Calosphaeriales. Mycologia
77(4):549–565
Barr ME (1990) Prodromus to nonlichenized, pyrenomycetous mem-
bers of class Hymenomycetes. Mycotaxon 39:43–184
Barr ME (1993) Redisposition of some taxa described by J. B. Ellis.
Mycotaxon 46:45–76
Barr ME (1994) Notes on the Amphisphaeriaceae and related families.
Mycotaxon 51:191–224
Barr ME, Rogers JD, Ju YM (1993) Revisionary studies in the Calo-
sphaeriales. Mycotaxon 48:529–535
Barrasa JM, Solans MJ, Moreno G (1985) Strattonia dissimilis (Sor-
dariales), una nueva especie coprófila. Int J Mycol Lichenol
2(1):75–84
Bartoli A, Maggi O, Persiani AM (1984) Effetia, a new genus of asco-
mycetes from tropical forest soils. Mycotaxon 19:515–522
Bates ST, Miller AN, Collections TM, Consortia MC (2018) The pro-
tochecklist of North American nonlichenized Fungi. Mycologia
110(6):1222–1348
Batsch AJGK (1786) Elenchus fungorum. Continuatio prima, pp
1–279
Bell A (2000) Podospora petrogale Fungi: Sordariales: Lasiospha-
eriaceae, a new species from Australia. Mueller 12(2):235–240
Bell A (2005) An illustrated guide to the coprophilous Ascomycetes of
Australia. CBS Biodivers Ser 3:1–172
Bell A, Mahoney DP (1997) Coprophilous fungi in New Zealand.
II. Podospora species with coriaceous perithecia. Mycologia
89(6):908–915
Bellemère A (1994) Asci and ascospores in ascomycete systematics.
In: Hawksworth DL (ed) Ascomycete systematics: problems
Fig. 69 Tulipispora ingoldii:
(redrawn from Révay etal.
2009). a Developing conidia
showing the separating cells. b
Main axes and conidia bearing
a secondary branch. c Conidio-
phores with conidia. Scale bars:
a–c = 20µm
Fungal Diversity
1 3
and perspectives in the nineties. Plenum Press, New York, pp
111–126
Berbee ML, Taylor JW (1992) Detecting morphological convergence
in true fungi, using 18S rRNA gene sequence data. Biosystems
28:117–125
Berlese AN (1900) Icones Fungorum Pyrenomycetes Sphaeriaceae.
Allantosporae 3(1–2):1–52
Boedijn KB (1934) Über die neue Gattung Trigonia van Beyma thoe
Kingma. Annales Mycologici 32(3–4):302–302
Boedijn KB (1962) The Sordariaceae of Indonesia. Persoonia
2(3):305–320
Boudier E (1904) Sur un nouveau genre et une nouvelle espèce de
Myriangiacées, le Guilliermondia saccoboloides. Bulletin De
La Société Mycologique De France 20:19–22
Bundhun D, Maharachchikumbura SSN, Jeewon R etal (2020) https://
sordariomycetes.org/, a platform for the identification, ranking.
Asian J Mycol 3(1):13–21
Cai L, Jeewon R, Hyde KD (2005) Phylogenetic evaluation and taxo-
nomic revision of Schizothecium based on ribosomal DNA and
protein coding genes. Fungal Divers 19:1–21
Cai L, Jeewon R, Hyde KD (2006a) Molecular systematics of Zopfiella
and allied genera: evidence from multi-gene sequence analyses.
Mycol Res 110(4):359–368
Cai L, Jeewon R, Hyde KD (2006b) Phylogenetic investigations of
Sordariaceae based on multiple gene sequences and morphology.
Mycol Res 110(2):137–150
Cain RF (1934) Studies of Coprophilous Spaeriales in Ontario, pp
1–126
Cain RF (1935) The genus Zygospermum. Mycologia 27(2):227
Cain RF (1948) Notes on seed-borne fungi. VI. Sordaria. Can J Res
26(5):486–495
Cain RF (1957a) Studies on coprophilous Ascomycetes. VI. Species
from the Hudson Bay area. Can J Bot 35(3):255–268
Cain RF (1957b) Synaptospora, a new genus of amerosporous Ascohy-
meniales (Ascomycetes). Beihefte Zur Sydowia 1:4–8
Cain RF (1961) Anixiella and Diplogelasinospora, two genera with
cleistothecia and pitted ascospores. Can J Bot 39(7):1667–1677
Cain RF, Mirza JH (1969) Apodospora, a new genus of the Sordari-
aceae. Can J Bot 4(5):891–896
Cain RF, Mirza JH (1972) Three new species of Arnium. Can J Bot
50:333–336
Calduch M, Gené J, Stchigel AM etal (2004) Ramophialophora, a new
anamorphic genus of Sordariales. Stud Mycol 50:83–88
Carmarán CC, Berretta M, Martínez S etal (2015) Species diversity
of Cladorrhinum in Argentina and description of a new species,
Cladorrhinum Australe. Mycol Prog 14(94):1–11
Carroll GC, Munk A (1964) Studies on lignicolous Sordariaceae.
Mycologia 56:77–98
Cesati V (1854) Herbarium Vivum Mycologicum 19: no. 1859
Cesati V (1856) Explicatio Iconum. Hedwigia 1(15):103–104
Cesati V, de Notaris G (1863) Schema di classificazione degle sferia-
cei italici aschigeri piu’ o meno appartenenti al genere Sphaeria
nell’antico significato attribuitoglide Persono. Commentario
Della Società Crittogamologica Italiana 1(4):177–420
Chang J-H, Kao H-W, Wang Y-Z (2010) Molecular phylogeny of Cer-
cophora, Podospora, Schizothecium (Lasiosphaeriaceae, Pyreno-
mycetes). Taiwania 55:110–116
Chernomor O, von Haeseler A, Minh BQ (2016) Terrace aware data
structure for phylogenomic inference from supermatrices. Syst
Biol 65:997–1008
Chesters CGC (1935) Studies on British pyrenomycetes: I. The life
histories of three species of Cephalotheca Fuck. Trans Br Mycol
Soc 19(4):261–279
Ciferri R (1954) Schedae Mycologicae XII-XXXIV. Sydowia
8(1–6):245–270
Clements FE (1909) The genera of Fungi, pp 1–227
Clements FE, Shear CL (1931) The genera of fungi. H.W. Wilson Co.,
New York
Cooke MC (1871) Handbook of British fungi. 2:489–981
Corda ACJ (1838) Icones fungorum hucusque cognitorum. 2:1–43
Crouan PL, Crouan HM (1867) Florule de Finistère, Contenant des
Descriptions de 360 Espèces Nouvelles de Sporogames. des
Nombreuses Observations, pp 1–262
Crous P, Shivas R, Quaedvlieg W etal (2014) Fungal planet description
sheets: 214–280. Persoonia 32:184–306
Currey F (1859) Synopsis of the fructification of the simple Sphaeriae
of the Hookerian herbarium. Trans Linn Soc Lond 22:313–355
Damm U, Crous PW, Fourie PH (2008) A fissitunicate ascus mecha-
nism in the Calosphaeriaceae, with novel species of Jattaea and
Calosphaeria on Prunus wood. Persoonia 20:39–52
Darriba D, Taboada GL, Doallo R etal (2012) jModelTest 2: more
models, new heuristics and parallel computing. Nat Methods
9:772–772
Davison EM, Davison PJ, Brims MH (2008) Moist chamber and field
collections of myxomycetes from the northern Simpson Desert,
Australia. Australas Mycol 27(3):129–135
De Notaris G (1867) Nuove reclute per la pirenomicetologia ital-
ica. Commentario Della Società Crittogamologica Italiana
2(3):477–492
Declercq B (2009) Omtrent Lasiosphaeria s.l. Sterbeeckia 28:35–41
Declercq B (2013) About Lasiosphaeria s.l. Sterbeeckia 35:3–5
del Valle CM, Romero AI (2003) Two new species of Camarops
(Boliniaceae, Ascomycotina) and a key to Argentinean species.
Sydowia 57(1):3–18
del Valle CM, Romero AI, Huhndorf SM etal (2011) A new spe-
cies and new records of Cercophora from Argentina. Mycologia
103(6):1372–1383
Dennis RWG (1978) British Ascomycetes, 2nd edn. Cramer, Vaduz
Desmazières JBHJ (1837) Notice sur quelques plantes cryptogames
nouvellement découvertes, en France. Annales Des Sciences
Naturelles Botanique 8:5–11
Desmazières JBHJ (1849) Dix-septième notice sur les plantes crypto-
games récemment découvertes en France. Annales Des Sciences
Naturelles Botanique Sér 3(11):339–365
Dettman JR, Harbinski FM, Taylor JW (2001) Ascospore morphology
is a poor predictor of the phylogenetic relationships of Neuros-
pora and Gelasinospora. Fungal Genet Biol 34(1):49–61
Dissanayake AJ, Bhunjun CS, Maharachchikumbura SSN etal (2020)
Applied aspects of methods to infer phylogenetic relationships
amongst fungi. Mycosphere 11(1):2652–2676
Doidge EM (1950) The South African fungi and lichens. Pretoria,
South Africa. Bothalia 5:1–1094
Doveri F (2008) A bibliography of Podospora and Schizothecium, a key
to the species, and a description of Podospora dasypogon newly
recorded from Italy. Pagine Di Micologia 29:61–159
Doveri F (2010) Une nouvelle variété de Tripterosporella sur crottin
- Une occasion de recombiner le pyrénomycète cléistothécial
Cercophora heterospora dans le genre Tripterosporella. Bulletin
Mycologique Et Botanique Dauphiné-Savoie 196:49–55
Dowding ES (1933) Gelasinospora, a new genus of pyrenomycetes
with pitted spores. Can J Res 9:294–305
Dülger B, Akata I (2016) Lasiosphaeria ovina, the first record for fam-
ily Lasiosphaeriaceae in Turkey. Nisan 7(1):88–91
Ekanayaka AH, Hyde KD, Gentekaki E etal (2019) Preliminary clas-
sification of leotiomycetes. Mycosphere 10(1):310–489. https://
doi. org/ 10. 5943/ mycos phere/ 10/1/7
Eliasson U (1971) Swedish find of Fragosphaeria purpurea Shear, a
cleistothecial ascomycete. Svensk Bot Tidskr 65:351–354
Ellis JB, Everhart BM (1892) The North American pyrenomycetes. A
contribution to mycologic botany
Ellis JB, Everhart BM (1893) New west American fungi. Erythea
1:197–206
Fungal Diversity
1 3
Ellis JB, Everhart BM (1895) New species of fungi from various locali-
ties. Proc Acad Natl Sci Phila 47:413–441
Eriksson OE, Hawksworth DL (1993) Outline of the Ascomy-
cetes–1993. Systema Ascomycetum 12:51–257
Eriksson OE, Winka K (1997) Supraordinal taxa of Ascomycota.
Myconet 1(1):1–16
Fakirova (1973) In: Dokl. Bolgarskoi Akademii Nauk, Biol 26(4):526
Fernández FA, Miller AN, Huhndorf SM etal (2006) Systematics of
the genus Chaetosphaeria and its allied genera: morphological
and phylogenetic diversity in north temperate and neotropical
taxa. Mycologia 98(1):121–130
Forin N, Vizzini A, Fainelli F, Ercole E, Baldan B (2021) Taxonomic
re-examination of nine Rosellinia types (Ascomycota, Xylariales)
stored in the Saccardo Mycological Collection. Microorganisms
9(3):666
Fort F, Cano J, Guarro J etal (1990) Contribución al estudio de
los ascomicetos del suelo de España. XI. Boletín Sociedad
Micológica De Madrid 14:61–73
Francuz B, Yera H, Geraut L etal (2010) Occupational asthma induced
by Chrysonilia sitophila in a worker exposed to coffee grounds.
Clin Vaccine Immunol 17(10):1645–1646
Frederick L, Uecker FA, Benjamin CR (1969) A new species of Neu-
rospora from soil of west Pakistan. Mycologia 61:1077–1084
Fries EM (1816) Uppstållning af de i Sverige funne Vårtsvampar (Scle-
romyci) 3:140–141
Fries EM (1823a) Sistens fungorum ordines, genera et species, huc
usque cognitas, quas ad normam methodi naturalis determinavit.
Systema Mycologicum 2:1–620
Fries EM (1823b) Synopsis Scleromycetum in Suecia nuper detecto-
rum, praemissa nova Sphaeriarum dispositione. Mykologische
Hefte 2:31–60
Fries EM (1828) Elenchus Fungorum. 2:1–154
Fries EM (1849) Summa vegetabilium Scandinaviae 2:259–572
Fuckel L (1870) Symbolae mycologicae. Beiträge zur Kenntniss der
Rheinischen Pilze. Jahrbücher Des Nassauischen Vereins Für
Naturkunde 23–24:1–459
Fuckel L (1872) Symbolae mycologicae. Beiträge zur Kenntniss der
rheinischen Pilze. Erster Nachtrag. Jahrbücher Des Nassauischen
Vereins Für Naturkunde 25–26:287–346
García D, Stchigel AM, José C etal (2004) A synopsis and re-
circumscription of Neurospora (syn. Gelasinospora) based
on ultrastructural and 28S rDNA sequence data. Mycol Res
108(10):1119–1142
Goh TK, Hyde KD (1998) A synopsis of and a key to Diplococcium
species, based on the literature, with a description of a new spe-
cies. Fungal Divers 1:65–83
Goh TK, Hyde KD, Umali TE (1998) Two new species of Diplococ-
cium from the tropics. Mycologia 90:514–517
Górz A, Boroń P (2018) Episternus onthophagi: a new monotypic
genus of epizoic fungus found on Onthophagus beetles (Scara-
baeoidea). Phytotaxa 376(1):43–59
Gramaje D, Mostert L, Groenewald JZ etal (2015) Phaeoacremo-
nium: from esca disease to phaeohyphomycosis. Fungal Biol
119(9):759–783
Grognet P, Silar P (2015) Maintaining heterokaryosis in pseudo-
homothallic fungi. Commun Integr Boil 8(4):e994382
Guarro J, Cano J (1988) The genus Triangularia. Trans Br Mycol Soc
91(4):587–591
Guarro J, von Arx JA (1987) The ascomycete genus Sordaria. Persoo-
nia 13:301–313
Guarro J, Cannon PF, van der Aa HA (1991) A synopsis of the genus
Zopfiella (Ascomycetes, Lasiosphaeriaceae). Systema Ascomy-
cetum 10(2):79–112
Hall TA (1999) BioEdit: a user-friendly biological sequence alignment
editor and analysis program for Windows 95/98/NT. Nucleic
Acids Symp Ser 41:95–98
Hawksworth DL, Eriksson OE (1986) The names of accepted orders of
Ascomycetes. Systema Ascomycetum 5:175–184
Hawksworth DL, Manoharachary C (1978) Isia, a new genus in the
Sordariaceae sensu lato for Thielavia neocaledoniensis. Trans
Br Mycol Soc 71(2):332–335
Hennebert GL (1968) Echinobotryum, Wardomyces and Mammaria.
Trans Br Mycol Soc 51(5):749–762
Hennings P (1898) Die in den Gewächshäusern des Berliner Botanis-
chen Gartens beobachteten Pilze. Verhandlungen Des Botanis-
chen Vereins Der Provinz Brandenburg 40:109–177
Hernández-Restrepo M, Gené J, Castañeda-Ruiz RF etal (2017)
Phylogeny of saprobic microfungi from Southern Europe. Stud
Mycol 86:53–97
Hilber O, Hilber R (1979) Einige Anmerkungen zu der Gattung Cer-
cophora Fuckel (Lasiosphaeriaceae). Zeitschrift Für Mykologie
45(2):209–233
Hilber R, Hilber O (2002) The Genus Lasiosphaeria and allied taxa,
pp 1–9
Hirooka Y, Rossman AY, Samuels GJ etal (2012) A monograph of
Allantonectria, Nectria, and Pleonectria (Nectriaceae, Hypo-
creales, Ascomycota) and their pycnidial, sporodochial, and
synnematous anamorphs. Stud Mycol 71:1–210
Hoffmann GF (1790) Vegetabilia cryptogama, vol 2
Holec J (2005) Distribution and ecology of Camarops tubulina (Asco-
mycetes, Boliniaceae) in the Czech Republic and remarks on its
European distribution. Czech Mycol 57(1/2):97
Holm L (1975) Nomenclatural notes on pyrenomycetes. Taxon
24:475–488
Hongsanan S, Sánchez-Ramírez S, Crous PW etal (2016) The evolu-
tion of fungal epiphytes. Mycosphere 7(11):1690–1712
Hongsanan S, Maharachchikumbura SS, Hyde KD etal (2017) An
updated phylogeny of Sordariomycetes based on phylogenetic
and molecular clock evidence. Fungal Divers 84:25–41
Hu DM, Cai L, Hyde KD etal (2006) The genera Podospora and Schi-
zothecium from mainland China. Cryptogam, Mycol 27(2):1–22
Huang S-K, Jeewon R, Hyde KD etal (2018) Beta-tubulin and Actin
gene phylogeny supports Phaeoacremonium ovale as a new spe-
cies from freshwater habitats in China. MycoKeys 41:1–15
Hughes SJ (1979) Relocation of species of Endophragmia auct. With
notes on relevant generic names. NZ J Bot 17:139–188
Huhndorf SM, Miller AN (2008) A new species of Camarops and
phylogenetic analysis of related taxa in the Boliniaceae. N Am
Fungi 3:231–239
Huhndorf SM, Fernández DJ, Lodge DJ (1999a) Noetropical Asco-
mycetes 9. Jobellisia species from Puerto Rico and elsewhere.
Sydowia 51(2):183–196
Huhndorf SM, Fernández F, Candoussau F (1999b) Two new species
of Synaptospora. Sydowia 51:176–182
Huhndorf SM, Miller AN, Fernández FA (2004a) Molecular systemat-
ics of the Coronophorales and new species of Bertia, Lasiobertia
and Nitschkia. Mycol Res 108(12):1384–1398
Huhndorf SM, Miller AN, Fernández FA (2004b) Molecular systemat-
ics of the Sordariales: the order and the family Lasiosphaeriaceae
redefined. Mycologia 96(2):368–387
Huhndorf SM, Miller AN, Fernández FA etal (2005) Neotropical
Ascomycetes 13. Cornipulvina and Erythromada, two new gen-
era from the Caribbean and elsewhere. Fungal Divers 20:59–69
Hyde KD (1994) Fungi from palms. X. Lockerbia palmicola, a new
cleistothecial genus in the Sordariales. Sydowia 46:23–28
Hyde KD, Goh TK (1999) Fungi on submerged wood from the River
Coln, England. Mycol Res 103(12):1561–1574
Hyde KD, Wong SW (2000) Annulatascus fusiformis sp. nov., a
new freshwater ascomycete from the Philippines. Mycologia
92:353–557
Fungal Diversity
1 3
Hyde KD, Frölich J, Taylor JE (1998) Fungi from palms XXVI. Reflec-
tions on unitunicate ascomycetes with apiospores. Sydowia
50:21–80
Hyde KD, Jones EBG, Liu JK etal (2013) Families of Dothideomy-
cetes. Fungal Divers 63:1–313
Hyde KD, Maharachchikumbura SS, Hongsanan S etal (2017) The
ranking of fungi: a tribute to David L. Hawksworth on his 70th
birthday. Fungal Divers 84(1):1–23
Hyde KD, Norphanphoun C, Maharachchikumbura SSN et al
(2020) Refined families of Sordariomycetes. Mycosphere
11(1):305–1059
Hyde KD, Bao DF, Hongsanan S etal (2021) Evolution of freshwater
Diaporthomycetidae (Sordariomycetes) provides evidence for
five new orders and six new families. Fungal Divers 14:71–105
Index Fungorum (2020) http:// www. index fungo rum. org/- N ame/ Names.
asp
Ito T, Nakagiri A (1994) Stellatospora, a new genus of the Sordari-
aceae. Mycoscience 35:413–415
Jacobson DJ, Powell AJ, Dettman JR etal (2004) Neurospora in tem-
perate forests of western North America. Mycologia 96:66–74
Jayawardena RS, McKenzie EHC, Chen YJ etal (2019) A database to
enhance identification of phytopathogenic genera. Asian J Mycol
2(1):281–286
Jeng RS, Krug JC (1976) Emblemospora, a new genus of the Sordari-
aceae. Can J Bot 54(16):1971–1976
Jeng RS, Krug JC (1977) New records and new species of Arnium.
Can J Bot 55:83–95
Jensen CN (1912) Fungous flora of the soil. Bull Cornell Univ Agric
Exp Stn 315:415–501
Jones EBG, Sakayaroj J, Suetrong S etal (2009) Classification of
marine Ascomycota, anamorphic taxa and Basidiomycota. Fun-
gal Divers 35:1–187
Jones EBG, Suetrong S, Sakayaroj J etal (2015) Classification of
marine Ascomycota, Basidiomycota, Blastocladiomycota and
Chytridiomycota. Fungal Divers 73:1–72
Jones EBG, Devadatha B, Abdel-Wahab MA etal (2020) Phylogeny of
new marine Dothideomycetes and Sordariomycetes from man-
groves and deep-sea sediments. Bot Mar 63(2):155–181
Ju Y-M, Rogers JD, Huhndorf SM (1996) Valsaria and notes on Endox-
ylina, Pseudothyridaria, Pseudovalsaria and Roussoella. Myco-
taxon 58:419–481
Kalyaanamoorthy S, Minh BQ, Wong TFK etal (2017) ModelFinder:
fast model selection for accurate phylogenetic estimates. Nat
Methods 14:587–589
Karsten PA (1873) Mycologia fennica. Pars secunda. Pyrenomycetes.
Bidrag till Kännedom Av Finlands Natur Och Folk 23:1–252
Katoh K, Standley DM (2013) MAFFT: iterative refinement and addi-
tional methods. Methods Mol Biol 1079:131–146
Khan RS, Krug JC (1989) New records of the Sordariaceae from East
Africa. Mycologia 81:862–869
Khan RS, Krug JC (1991) Podospora fibrinocaudata, a new species
from California. Mycologia 83(6):817–821
Kirk PM, Cannon PF, David JC etal (2001) Ainsworth and Bisby’s
dictionary of the fungi, 9th edn. CABI International, Egham,
pp 1–655
Kirk PM, Cannon PF, Minter DW etal (2008) Ainsworth and Bisby’s
dictionary of the fungi, 10th edn. CAB International, Wallingford
Kondratyuk S, Lőkös L, Tchabanenko S etal (2013) New and note-
worthy lichen-forming and lichenicolous fungi. Acta Biol Hung
55(3–4):275–349
Konta S, Hongsanan S, Eungwanichayapant PD etal (2017) Lepto-
sporella (Leptosporellaceae fam. nov.) and Linocarpon and
Neolinocarpon (Linocarpaceae fam. nov.) are accommodated in
Chaetosphaeriales. Mycosphere 8:1943–1974
Krug JC (1989) Periamphispora, a new genus of the Sordariaceae.
Mycologia 81:475–479
Krug JC (1995) The genus Fimetariella. Can J Bot 73(12):1905–1916
Krug JC, Cain FR (1972) Additions to the genus Arnium. Can J Bot
50:367–373
Krug JC, Jeng RS (1977) The genus Camptosphaeria. Sydowia
29(1–6):71–74
Krug JC, Khan RS (1991) A new homothallic species of Neurospora
from Hungary. Mycologia 83(6):829–832
Krug JC, Khan RS, Jeng RS (1994) A new species of Gelasinospora
with multiple germ pores. Mycologia 86:250–253
Krug JC, Scott JA (1994) The genus Bombardioidea. Can J Bot
72(9):1302–1310
Krug JC, Udagawa S, Jeng RS (1983) The genus Apiosordaria. Myco-
taxon 17:533–549
Kruys Å, Huhndorf SM, Miller AN (2014) Coprophilous contributions
to the phylogeny of Lasiosphaeriaceae and allied taxa within
Sordariales (Ascomycota, Fungi). Fungal Divers 70:101–113
Kummer V, Richter T, Schwik J (2005) Wegelina grumsiniana comb.
nov. (Ascomycetes, Calosphaeriales)-ein Pyrenomycet auf der
Porenschicht faulender Zunderschwämme (Fomes fomentarius).
Z Mykol 71:227–236
Lecocq T, Vereecken NJ, Michez D etal (2013) Patterns of genetic
and reproductive traits differentiation in mainland vs. Corsican
populations of bumblebees. PLoS ONE 8(6):e65642
Leroy P (2006) Jobellisia saliciluticola (Ascomycota - Sordariomyceti-
dae), une espèce nouvelle récoltée dans deux régions de France.
Documents Mycologiques 34:133–134
Liu F, Hu DM, Cai L (2012) Conlarium duplumascospora gen. et.
sp. nov. and Jobellisia guangdongensis sp. nov. from freshwater
habitats in China. Mycologia 104(5):1178–1186
Lloyd CG (1923) Mycological notes no. 68. Mycol Writ 7:1169–1184
Lodhi SA, Mirza F (1962) A new genus of the Eurotiales. Mycologia
54(2):217–219. https:// doi. org/ 10. 1080/ 00275 514. 1962. 12024
993
Lumbsch HT (2010) Myconet volume 14. Part one. Outline of Ascomy-
cota—2009. Part two. Notes on Ascomycete Systematics. Nos.
4751–5113. Fieldiana Life Earth Sci 1:1–64
Lundqvist N (1964a) Anopodium, a new genus of coprophilous
pyrenomycetes with apically pedicellate spores. Bot Notiser
117(4):355–365
Lundqvist N (1964b) Fimetariella, a new genus of coprophilous pyr-
enomycetes. Bot Notiser 117:238–248
Lundqvist N (1967) On spore germination in the Sordiaceae, exempli-
fied by the new cleistocarpous genus Copromyces. Arkiv Før
Botanik 6(7):327–337
Lundqvist N (1969) Zygopleurage and Zygospermella. Bot Notiser
122:353–374
Lundqvist N (1972) Nordic Sordariaceae s. lat. Symbolae Botanicae
Upsalliensis 20:1–374
Lundqvist N (1974) Studia fungorum fimi II. New records of Arnia,
and a newspecies, A. bellum. Svensk Bot Tidskr 68:289–303
Luo ZL, Hyde KD, Liu JK etal (2019) Freshwater Sordariomycetes.
Fungal Divers 99:451–660
Ma J, Zhang K, Zhang XG etal (2016) Three new species of Spa-
dicoides from Lushan Mountain, China. Mycol Prog 15(5):43
Madrid H, Cano J, Stchigel A etal (2010) Ramophialophora humi-
cola and Fibulochlamys chilensis, two new microfungi from soil.
Mycologia 102(3):605–612
Maharachchikumbura SS, Hyde KD, Jones EG etal (2015) Towards
a natural classification and backbone tree for Sordariomycetes.
Fungal Divers 72:199–301
Maharachchikumbura SS, Hyde KD, Jones EG etal (2016) Families
of sordariomycetes. Fungal Divers 79:1–317
Mahoney DP, Huang LH, Backus MP (1969) New homothallic Neuro-
sporas from tropical soils. Mycologia 61:264–274
Malloch D, Cain RF (1971) New cleistothecial Sordariaceae and a new
family, Coniochaetaceae. Can J Bot 49(6):869–880
Fungal Diversity
1 3
Marchal E (1885) Champignons coprophiles de Belgique. Bulletin de
la Société Royale de Botanique de Belgique 24:57–77
Marin-Felix Y, Miller AN, Cano-Lira JF etal (2020) Re-evaluation of
the order Sordariales: Delimitation of Lasiosphaeriaceae s. str.,
and introduction of the new families Diplogelasinosporaceae,
Naviculisporaceae, and Schizotheciaceae. Microorganisms
8(9):1430
Matsushima T (1975) Icones Microfungorum a Matsushima Lectorum.
T. Matsushima, Kobe
Matzer M (1993) Beitrag zur Kenntnis der Ascomycetengattungen
Globosphaeria, Roselliniopsis und Synaptospora. Cryptogam,
Mycol 14(1):11–19
Matzer M, Hafellner J (1990) Eine Revision der lichenicolen Arten der
Sammelgattung Rosellinia (Ascomycetes). Bibliotheca Licheno-
logica 37:1–138
Mercuri OA (1972) Camarops (Ascomycetes), género nuevo para la
Argentina. Darwiniana 17:548–551
Miller JH (1930) British Xylariaceae I. Trans Br Mycol Soc
15(1–2):134–154
Miller AN, Huhndorf SM (2004a) A natural classification of Lasio-
sphaeria based on nuclear LSU rDNA sequences. Mycol Res
108(1):26–34. https:// doi. org/ 10. 1017/ S0953 75620 30088 64
Miller AN, Huhndorf SM (2004b) Using phylogenetic species recogni-
tion to delimit species boundaries within Lasiosphaeria. Myco-
logia 96(5):1106–1127
Miller AN, Huhndorf SM (2005) Multi-gene phylogenies indicate
ascomal wall morphology is a better predictor of phylogenetic
relationships than ascospore morphology in the Sordariales
(Ascomycota, Fungi). Mol Phylogenet Evol 35(1):60–75
Miller AN, Huhndorf SM, Fournier J (2014) Phylogenetic relationships
of five uncommon species of Lasiosphaeria and three new spe-
cies in the Helminthosphaeriaceae (Sordariomycetes). Mycologia
106(3):505–524
Mirza JH, Cain RF (1969) Revision of the genus Podospora. Can J Bot
47(12):1999–2048
Mirza JH, Nasir MA (1968) Additions to the coprophilous fungi of
West Pakistan II. Nova Hedwigia 16:283–288
Moreau C (1953) Les genres Sordaria et Pleurage. Encyclopédie
Mycologique 25:1–330
Mostert L, Groenewald JZ, Summerbell RC etal (2006) Taxonomy and
pathology of Togninia (Diaporthales) and its Phaeoacremonium
anamorphs. Stud Mycol 54:1–113
Mouchacca J, Gams W (1993) The hyphomycete genus Cladorrhinum
and its teleomorph connections. Mycotaxon 48:415–440
Mouton V (1897) Troisième notice sur des ascomycètes nouveaux ou
peu connus. Bulletin De La Société Royale De Botanique De
Belgique 36:10–21
Mugambi GK, Huhndorf SM (2010) Multigene phylogeny of the Coro-
nophorales: morphology and new species in the order. Mycologia
102(1):185–210
Muggia L, Fleischhacker A, Kopun T, Grube M (2015) Extremotoler-
ant fungi from alpine rock lichens and their phylogenetic rela-
tionships. Fungal Divers 76:119–142
Müller E, von Arx JA (1962) Die Gattungen der didymosporen Pyreno-
myceten. Beitr. Kryptogamenflora Schweiz 11(2):1–922
Mungai PG, Chukeatirote E, Njogu JG etal (2012) Studies of copro-
philous ascomycetes in Kenya: Sordariales from wildlife dung.
Mycosphere 3(4):437–448
Munk A (1957) Danish Pyrenomycetes. A preliminary flora. Dansk
Botanisk Arkiv 17:1–491
Muroi T, Udagawa S-I, Otani Y (1987) Some coprophilous ascomy-
cetes from Peru. In: Inoue H (ed) Studies on cryptogams in
Southern Peru. Tokai University Press, Tokyo
Najwa AM, Sarah AA, Ahmed HFG etal (2012) Pleurostomophora
ochracea, a novel agent of human eumycetoma with yellow
grains. J Clin Microbiol 50:2987–2994
Nannfeldt JA (1932) Studien über die Morphologie und Systematik
der nichtlichenisierten inoperculaten Discomyceten. Nova Acta
Regiae Societatis Scientiarum Upsaliensis 8(2):1–368
Nannfeldt JA (1972) Camarops Karst (Sphaeriales–Boliniaceae).
Svensk Bot Tidskr 66:335–376
Negreiros AMP, Júnior RS, Rodrigues APMS etal (2019) Prevalent
weeds collected from cucurbit fields in Northeastern Brazil
reveal new species diversity in the genus Monosporascus. Ann
Appl Biol 174(3):349–363
Nguyen L-T, Schmidt HA, von Haeseler A etal (2015) IQ-TREE: a
fast and effective stochastic algorithm for estimating maximum
likelihood phylogenies. Mol Biol Evol 32:268–274
Niessl G (1872) Beiträge zur Kenntniss der Pilze. Beschreibung
neuer und wenig bekannter Pilze. Verhandlungen Des Naturfor-
schenden Vereins in Brünn 10:153–217
Niessl G (1883) Über die Theilung der Gattung Sordaria. Hedwigia
22:153–156
Nitschke TH (1867) Pyrenomycetes Germanici. Breslau
Nitschke TH (1873) Flora oder Botanische Zeitung 31:450
Parguey-Leduc A (1960) Étude des asques et du développement de 1’
Helminthosphaeria clavariarum (Desm.) Fuck. ap. Munk. Bul-
letin de la Société Mycologique de France 76:15–33
Perkins DD, Raju NB (1986) Neurospora discreta, a new heterothallic
species defined by its crossing behavior. Exp Mycol 10:323–338
Perkins DD, Turner BC (1988) Neurospora from natural populations:
toward the population biology of a haploid eukaryote. Exp Mycol
12:91–131
Perkins DD, Turner BC, Barry EG (1976) Strains of Neurospora col-
lected from nature. Evolution 30:281–313
Persoon C (1794) Dispositio methodica fungorum. Neues Magazin für
die Botanik 1:81–128
Persoon CH (1796) Observationes mycologicae, vol. 1
Persoon CH (1801) Synopsis methodica fungorum, pp 1–706
Petrak F (1923) Mykologische Notizen. VI. Annales Mycologici
21(3–4):182–335
Petrak F (1953) Fungi Beltsvillenses. Sydowia 7(1–4):121–132
Petrak F (1967) Ergebnisse einer Revision der Grundtypen ver-
schiedener Gattungen der Askomyzeten und Fungi imperfecti.
Sydowia 21(1–6):240–248
Pollack FG, Uecker FA (1974) Monosporascus cannonballus an unu-
sual ascomycete in cantaloupe roots. Mycologia 66:346–349
Potebnia A (1907) Mycologische Studien. Annales Mycologici
5(1):1–28
Rabenhorst GL (1851) Klotzschii herbarium vivum mycologicum
sistens fungorum per totam Germaniam crescentium collectio-
nem perfectam. Editio prima Centuria XVI:no. 1501–no. 1600
Rabenhorst G (1860) Fungi europaei exsiccati. Edito nova Series
secunda Cent 3:no. 250
Raja HA, Shearer CA (2008) Freshwater ascomycetes: new and note-
worthy species from aquatic habitats in Florida. Mycologia
100(3):467–489
Raja H, Schoch CL, Hustad V etal (2011) Testing the phylogenetic
utility of MCM7 in the Ascomycota. MycoKeys 1:63
Ramaley AW (2005) Sulcatistroma nolinae (Calosphaeriales), and its
Phialophora-like anamorph. Mycotaxon 93:139–144
Ranghoo VM, Tsui CKM, Hyde KD (2001) Brunneosporella aquatica
gen. et sp. nov., Aqualignicola hyalina gen. et sp. nov., Jobel-
lisia viridifusca sp. nov. and Porosphaerellopsis bipolaris sp.
nov. (ascomycetes) from submerged wood in freshwater habitats.
Mycol Res 105(5):625–633
Rappaz F (1995) Anthostomella and related Xylariaceous fungi on
hardwood from Europe and North America. Mycologia Helvetica
7:99–168
Rashmi M, Kushveer JS, Sarma VV (2019) A worldwide list of endo-
phytic fungi with notes on ecology and diversity. Mycosphere
10(1):798–1079
Fungal Diversity
1 3
Reagan DP, Waide RB (1996) The food web of a tropical rain forest,
vol 84. University of Chicago, Chicago
Réblová M (1998) Fungal diversity in the Czech Republic. New species
of Apiorhynchostoma, Capronia, Ceratosphaeria and Lasiospha-
eria. Sydowia 50:299–251
Réblová M (1999a) Studies in Chaetosphaeria sensu lato I. The gen-
era Chaetosphaerella and Tengiomyces gen. nov. of the Hel-
minthosphaeriaceae. Mycotaxon 70:387–420
Réblová M (1999b) Teleomorph-anamorph connections in Ascomy-
cetes 3: three new lignicolous species of Helminthosphaeria.
Sydowia 51:233–244
Réblová M (2002) Synaptospora olandica, a new species from Sweden.
Sydowia 54(2):248–255
Réblová M (2006) Molecular systematics of Ceratostomella sensu lato
and morphologically similar fungi. Mycologia 98:63–93
Réblová M (2007) Barbatosphaeria gen. et comb. nov., a new genus for
Calosphaeria barbirostris. Mycologia 99:723–732
Réblová M (2008) Bellojisia, a new sordariaceous genus for Jobel-
lisia rhynchostoma and a description of Jobellisiaceae fam. nov.
Mycologia 100(6):893–901
Réblová M (2011) New insights into the systematics and phylogeny
of the genus Jattaea and similar fungi of the Calosphaeriales.
Fungal Divers 49:167–198
Réblová M, Mostert L (2007) Romellia is congeneric with Togninia,
and description of Conidiotheca gen. nov. for one species of this
genus with polysporous asci. Mycol Res 111(3):299–307
Réblová M, Mostert L, Gams W etal (2004) New genera in the Calo-
sphaeriales: Togniniella and its anamorph Phaeocrella, and
Calosphaeriophora as anamorph of Calosphaeria. Stud Mycol
50(2):533–550
Réblová M, Gams W, Štěpánek V (2011) The new hyphomycete genera
Brachyalara and Infundichalara, the similar Exochalara and spe-
cies of Phialophoraʻ sect. Catenulataeʼ (Leotiomycetes). Fungal
Divers 46:67–86
Réblová M, Jaklitsch WM, Réblová K etal (2015) Phylogenetic recon-
struction of the Calosphaeriales and Togniniales using five genes
and predicted RNA secondary structures of ITS, and Flabel-
lascus tenuirostris gen. et sp. nov. PLoS ONE 10(12):e0144616
Réblová M, Miller AN, Rossman AY etal (2016) Recommendations
for competing sexual-asexually typified generic names in Sor-
dariomycetes (except Diaporthales, Hypocreales, and Magna-
porthales). IMA Fungus 7(1):131–153
Réblová M, Miller AN, Réblová K etal (2018) Phylogenetic classi-
fication and generic delineation of Calyptosphaeria gen. nov.,
Lentomitella, Spadicoides and Torrentispora (Sordariomycetes).
Stud Mycol 89:1–62
Rehm H (1904) Ascomycetes Americae borealis. Ann Myc 2:175
Révay Á, Gönczöl J, Descals E (2009) Tulipispora ingoldii (Hypho-
mycetes) gen. et sp. nov. from submerged wood. Nova Hedwigia
88(1–2):41–48
Rick JE (1931) Monographia Boliniarum Riograndensium. Brotéria
Série Botânica 25(2):65–71
Rogers JD (1988) Apiocamarops cryptocellula, a new species from
Guyana. Mycologia 80(5):738–741
Rogers JD, Ju YM (2003) Occultitheca costaricensis gen. et sp. nov.
and Apiocamarops pulvinata sp. nov. from Costa Rica. Sydowia
55(2):359–364
Rogers JD, Stone J, Ju YM (1994a) Anthostomella formosa var.
abietis var. nov. and notes on Apiorhynchostoma. Mycologia
86(5):700–703
Rogers JD, Yu YM, Læssøe T (1994b) Neohypodiscus nom. nov. for
Hypodiscus. Mycologia 86(5):684–690
Romero AI, Samuels GJ (1991) Studies on xylophilous fungi from
Argentina. VI. Ascomycotina on Eucalyptus viminalis (Myrta-
ceae). Sydowia 43:31–41
Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic
inference under mixed models. Bioinformatics 19(12):1572–1574
Saccardo PA (1878) Fungi Veneti novi vel critici vel mycologiae Ven-
etae addendi. Series IX. Michelia 1:361–445
Saccardo PA (1881) Fungi Gallici lecti a cl. viris P. Brunaud, C.G. Gil-
let, Abb. Letendre, A. Malbranche, J. Therry vel editi in Myco-
theca Gallica cl. C. Roumeguèri, recensuit P.A. Saccardo. Series
III Michelia 2(7):302–371
Saccardo PA (1882) Sylloge Pyrenomycetum, Vol. I. Sylloge Fungo-
rum 1:1–768
Saccardo PA (1886) Sylloge Hyphomycetum. Sylloge Fungorum
4:1–807
Saccardo PA (1891) Supplementum Universale, Pars I. Agaricaceae-
Laboulbeniaceae. Sylloge Fungorum 9:1–1141
Saccardo PA (1906) Micromycetes americani novi. J Mycol 12:47–51
Samuels GJ, Rogers JD (1987) Camarops flava sp. nov., Apiocamarops
alba gen. et sp. nov., and notes on Camarops scleroderma and C.
ustulinoides. Mycotaxon 28(1):45–59
Samuels GJ, Candoussau F, Magni JF (1997) Fungicolous pyrenomy-
cetes 1. Helminthosphaeria and the new family Helminthospha-
eriaceae. Mycologia 89:141–155
Schaumann K (1972) Veröffentlichungen des Instituts für Meeres-
forschung in Bremerhaven, vol 14
Schoch CL, Robbertse B, Robert V etal (2014) Finding needles in
haystacks: linking scientific names, reference specimens and
molecular data for Fungi. Database
Schröter J (1894) Kryptogamen-Flora von Schlesien 3–2(9):257–384
Schröter J (1897) Kryptogamen-Flora von Schlesien 3–2(10):385–500
Senanayake IC, Crous P, Groenewald J etal (2017) Families of Dia-
porthales based on morphological and phylogenetic evidence.
Stud Mycol 86:217–296
Senanayake IC, Jeewon R, Chomnunti P etal (2018) Taxonomic cir-
cumscription of Diaporthales based on multigene phylogeny and
morphology. Fungal Divers 93:241–443
Senanayake IC, Rathnayaka AR, Marasinghe DS etal (2020) Mor-
phological approaches in studying fungi: collection, exami-
nation, isolation, sporulation and preservation. Mycosphere
11(1):2678–2754
Shaw DE (1998) Species of Neurospora recorded in Australia, and the
collection of Neurospora conidia by honey bees in lieu of pollen.
Mycologist 4(12):154–158
Shear CL (1923) Life histories and undescribed genera and species of
fungi. Mycologia 15(3):120–131
Shear CL (1938) Mycological notes. II. Mycologia 30:580–593
Shear CL (1940) Mycological notes. IV. Mycologia 32:541–549
Shear CL, Dodge BO (1927) Life histories and heterothallism of the
red bread-mold fungi of the Monilia sitophila group. J Agric Res
34(11):1019–1041
Shenoy BD, Jeewon R, Hyde KD (2007) Impact of DNA sequencedata
on the taxonomy of anamorphic fungi. Fungal Divers 26:1–54
Shenoy BD, Jeewon R, Wang HK etal (2010) Sequence data reveals
phylogenetic affinities of fungal anamorphs Bahusutrabeeja,
Diplococcium, Natarajania, Paliphora, Polyschema, Rattania
and Spadicoides. Fungal Divers 44:161–169
Sivanesan A (1983) Studies on ascomycetes. Trans Br Mycol Soc
81:313–332
Spatafora JW (1995) Ascomal evolution of filamentous ascomycetes:
evidence from molecular. Can J Bot 73:811–815
Spatafora JW, Blackwell M (1993) Molecular systematics of unitu-
nicate perithecial ascomycetes: the Clavicipitales-Hypocreales
connection. Mycologia 85:912–922
Spatafora JW, Blackwell M (1994) The polyphyletic origins of ophios-
tomatoid fungi. Mycol Res 98(1):1–9
Spatafora JW, Sung G-H, Johnson D etal (2006) A five-gene phylo-
geney of Pezizomycotina. Mycologia 98(6):1018–1028
Fungal Diversity
1 3
Spegazzini C (1888) Fungi Fuegiani. Boletín De La Academia
Nacional De Ciencias En Córdoba 11(2):135–311
Spooner BM (1986) New or rare British microfungi from Esher Com-
mon, Surrey. Trans Br Mycol Soc 86(3):401–408
Stadler M, Kuhnert E, Peršoh D etal (2013) The xylariaceae as model
example for a unified nomenclature following the BOne fungus–
one name (1F1N) concept. Mycology 4(1):5–21
Stafleu FA, Voss EG (1969) Synopsis of proposals on botanical nomen-
clature. Seattle 1969. Regnum Veget 60:1–124
Stchigel AM, Cano J, Miller AN etal (2006) Corylomyces: a new
genus of Sordariales from plant debris in France. Mycol Res
110(11):1361–1368
Subramanian CV, Lodha BC (1968) Two interesting coprophilous fungi
from India. Curr Sci 37:245–248
Subramanian CV, Sekar G (1986) Pseudocercophora ingoldii gen. et
sp. nov. and its Mammaria anamorph. J Singap Natl Acad Sci
15:58–60
Suh SO, Blackwell M (1999) Molecular phylogeny of the cleistothecial
fungi placed in Cephalothecaceae and Pseudeurotiaceae. Myco-
logia 91:836–848
Sun JZ, Liu XZ, McKenzie EHC etal (2019) Fungicolous fungi: termi-
nology, diversity, distribution, evolution, and species checklist.
Fungal Divers 95:337–430
Sutton BC (1973) Hyphomycetes from Manitoba and Saskatchewan,
Canada. Mycol Pap 132:1–143
Svrček M (1969) Bolinia tubulina (Alb. Et Schw. ex Fr.) Sacc. v
Československu [Bolinia tubulina (Alb. Et Schw. ex Fr.) Sacc.
in der Tschechoslowakei]. Čes Mykol 23:118–122
Tai FL (1935) Two new species of Neurospora. Mycologia 27:328–330
Tang AMC, Jeewon R, Hyde KD (2007) Phylogenetic utility of protein
(RPB2, β-tubulin) and ribosomal (LSU, SSU) gene sequences
in the systematics of Sordariomycetes (Ascomycota, Fungi).
Antonie Van Leeuwenhoek 91(4):327–349. https:// doi. org/ 10.
1007/ s10482- 006- 9120-8
Taylor JW, Natvig DO (1989) Mitochondrial DNA and evolution of
heterothallic and pseudohomothallic Neurospora species. Mycol
Res 93(3):257–272
Theissen F (1909) Xylariaceae Austro-Brasilienses. II. Ann Myc
7:1–18
Tode HJ (1791) Fungi Mecklenburgenses Selecti, vol 2
Trail F, Wang Z, Stefanko K etal (2017) The ancestral levels of tran-
scription and the evolution of sexual phenotypes in filamentous
fungi. PLoS Genet 13(7):e1006867
Tulasne LR, Tulasne C (1863) Selecta Fungorum Carpologia, Tomus
Secundus. Xylariei - Valsei - Sphaeriei 2:1–319
Turner BC, Perkins DD, Fairfield A (2001) Neurospora from natural
populations: a global study. Fungal Genet Biol 32:67–92
Udagawa S (1980) New or noteworthy Ascomycetes from southeast
Asian soil I. Trans Mycol Soc Jpn 21(1):17–34
Udagawa SI, Furuya K (1974) Notes on some Japanese Ascomycetes
XIII. Trans Mycol Soc Jpn 15(3):206–214
Udagawa S, Muroi T (1979) Copropilous Pyrenomycetes from Japan
V. Trans Mycol Soc Jpn 20:453–468
Udagawa S, Sugiy Y (1982) In: Reports on the cryptogamic study in
Nepal (Tokyo), p 25
Ueda S (1994) A new Cercophora with a Chrysosporium–like anamo-
rph. Mycoscience 35:287–290. https:// doi. org/ 10. 1007/ BF022
68451
Untereiner WA (1993) A taxonomic revision of the genus Endoxyla.
Mycologia 85(2):294–310
Untereiner WA, Bogale M, Carter A etal (2013) Molecular phylogeny
of Boliniales (Sordariomycetes) with an assessment of the sys-
tematics of Apiorhynchostoma, Endoxyla and Pseudovalsaria.
Mycologia 105(3):564–588
Van Beyma Thoe Kingma FH (1933) Beschreibung einiger neuer
Pilzarten aus dem Centraalbureau voor Schimmelcultures
II—Baarn (Holland). Zentralblatt Für Bakteriologie Und Para-
sitenkunde, Abteilung 2(89):236–243
Varghese KIM, Rao VG (1979) Two new additions to the Indian Asco-
mycetes. Biovigyanam 5:1–4
Vasilyeva LN (1997) Camarops pugillus (Schw.: Fr.) Shear from the
Russian far East. Mycol Phytopatol 31:5–7
Vasilyeva LN (2007) Pyrenomycetes of the Russian Far East 2. Mol-
licamarops stellata gen. et sp. nov. Mycotaxon 99:159–162
Vasilyeva LN, Stephenson SL, Miller AN (2007) Pyrenomycetes of
the Great Smoky Mountains National Park. IV. Biscogniauxia,
Camaropella, Camarops, Camillea, Peridoxylon and Whalleya.
Fungal Divers 25:219–231
Vijaykrishna D, Mostert L, Jeewon R etal (2004) Pleurostomophora,
an anamorph of Pleurostoma (Calosphaeriales), a new anamo-
rph genus morphologically similar to Phialophora. Stud Mycol
50:387–395
Vojtkova H, Simonovicova A, Cernansky S (2020) Neurospora sit-
ophila in indoor environment of buildings. In IOP conference
series: earth and environmental science 444(1):012055
von Arx JA (1975) On Thielavia and some similar genera of Ascomy-
cetes. Stud Mycol 8:1–31
von Arx JA (1981a) The genera of fungi sporulating in pure culture,
3re edn. Cramer, Vaduz, p 424
von Arx JA (1981b) On Monilia sitophila and some families of asco-
mycetes. Sydowia 34:13–29
von Arx JA (1982) A key to the species of Gelasinospora. Persoonia
11:443–449
von Arx JA, Müller E (1954) Die Gattungen der amerosporen Pyr-
enomyceten. Beiträge Zur Kryptogamenflora Der Schweiz
11(1):1–434
von Arx JA, Guarro J, van der Aa HA (1987) Asordaria, a new genus of
the Sordariaceae, and a new species of Melancarpous. Persoonia
13:263–272
von Höhnel F (1917) Mykologische Fragmente. Nrn. 120–190. Annales
Mycologici 15(5):293–383
Vu D, Groenewald M, De Vries M etal (2019) Large-scale generation
and analysis of filamentous fungal DNA barcodes boosts cover-
age for kingdom fungi and reveals thresholds for fungal species
and higher taxon delimitation. Stud Mycol 92:135–154
Walker DM, Castlebury LA, Rossman AY etal (2012) New molecular
markers for fungal phylogenetics: two genes for species-level
systematics in the Sordariomycetes (Ascomycota). Mol Phylo-
genet Evol 64(3):500–512
Wallroth CFW (1833) Flora Cryptogamica Germaniae. 2:1–923
Wang YZ, Aptroot A, Hyde KD (2004) Revision of the Ascomycete
genus Amphisphaeria. Fungal Divers Res Ser 13:1–168
Wang XW, Houbraken J, Groenewald JZ etal (2016) Diversity and tax-
onomy of Chaetomium and chaetomium-like fungi from indoor
environments. Stud Mycol 84:145–224
Wang XW, Bai FY, Bensch K etal (2019a) Phylogenetic re-eval-
uation of Thielavia with the introduction of a new family
Podosporaceae. Stud Mycol 93:155–252
Wang XW, Yang FY, Meijer M etal (2019b) Redefining Humicola
sensu stricto and related genera in the Chaetomiaceae. Stud
Mycol 93:65–153
Watanabe T (1989) Three species of Sordaria, and Eudarluca biconica
from cherry seeds. Trans Mycol Soc Jpn 30:395–400
Weitzman I, Silva-Hutner M (1967) Non-keratinous agar media as sub-
strates for the ascigerous state in certain members of the Gym-
noascaceae pathogenic for man and animals. Sabouraudia 5:335–340
Whitehouse HLK (1949) Heterothallism and sex in the fungi. Biol Rev
24(4):411–447
Wijayawardene NN, Hyde KD, Tibpromma S etal (2017) Towards
incorporating asexual fungi in a natural classification: checklist
and notes 2012–2016. Mycosphere 8(9):1457–1555
Fungal Diversity
1 3
Wijayawardene NN, Hyde KD, Lumbsch HT etal (2018) Outline of
ascomycota: 2017. Fungal Divers 88:167–263
Wijayawardene NN, Hyde KD, Al-Ani LKT etal (2020) Outline of
Fungi and fungus-like taxa. Mycosphere 11(1):1060–1456
Winter G (1884) Rabenhorst's Kryptogamen-Flora, Pilze - Ascomy-
ceten, vol 1. vol 2
Winter G (1885) Rabenhorst’s Kryptogamen-Flora. Pilze-Ascomy-
ceten, Edn 2 1(2):193–528
Winter G (1886) Ascomyceten: Gymnoasceen und Pyenomyceten. In:
Rabenhorst’s Kryptogamen-Flora. Eduard Kummer, Leipzig, vol
1, p 928
Yaguchi T, Sano A, Yarita K etal (2006) New species of Cephalotheca
isolated from a Korean patient. Mycotaxon 96:309–322
Zhang N, Blackwell M (2001) Molecular phylogeny of dogwood
anthracnose fungus (Discula destructiva) and the Diaporthales.
Mycologia 93(2):355–365
Zhang N, Castlebury LA, Miller AN etal (2006) An overview of the
systematics of the Sordariomycetes based on a four-gene phylog-
eny. Mycologia 98(6):1076–1087
Zhang ZF, Liu F, Zhou X etal (2017) Culturable mycobiota from Karst
caves in China, with descriptions of 20 new species. Persoonia:
Mol Phylogeny Evol Fungi 39:1–31
Zopf W (1896) Übersicht der auf Flechten schmarotzenden Pilze. Hed-
wigia 35:312–366
Authors and Aliations
Shi‑KeHuang1,2,3,4,5· KevinD.Hyde3,6· AusanaMapook3· SajeewaS.N.Maharachchikumbura7·
Jayarama D.Bhat8,9· EricH.C.McKenzie10· RajeshJeewon11· Ting‑ChiWen1,2,5
Shi-Ke Huang
cocohuangsk@gmail.com
Kevin D. Hyde
kdhyde3@gmail.com
Ausana Mapook
phung.ausana@gmail.com
Sajeewa S. N. Maharachchikumbura
sajeewa83@yahoo.com
D. Jayarama Bhat
bhatdj@gmail.com
Eric H. C. McKenzie
MckenzieE@landcareresearch.co.nz
Rajesh Jeewon
r.jeewon@uom.ac.mu
1 State Key Laboratory Breeding Base ofGreen Pesticide
andAgricultural Bioengineering, Key Laboratory ofGreen
Pesticide andAgricultural Bioengineering, Ministry
ofEducation, Guizhou University, Guiyang550025, China
2 The Engineering Research Center ofSouthwest
Bio-Pharmaceutical Resources, Ministry ofEducation, Guizhou
University, Guiyang550025, GuizhouProvince, China
3 Center ofExcellence inFungal Research, Mae Fah Luang
University, ChiangRai57100, Thailand
4 School ofScience, Mae Fah Luang University,
ChiangRai57100, Thailand
5 The Mushroom Research Centre, Guizhou University,
Guiyang550025, China
6 Institute ofPlant Health, Zhongkai University ofAgriculture
andEngineering, Haizhu District, Guangzhou510225, China
7 School ofLife Science andTechnology, University
ofElectronic Science andTechnology ofChina,
Chengdu611731, China
8 Department ofBotany, Goa University, Goa, India
9 Present Address: GoaVelha, India
10 Manaaki Whenua-Landcare Research, Private Mail Bag
92170, Auckland, NewZealand
11 Department ofHealth Sciences, Faculty ofMedicine
andHealth Sciences, University ofMauritius, Reduit,
Mauritius
