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Tubeufiales, ord. nov., integrating sexual and asexual
generic names
Saranyaphat Boonmee & Amy Y. Rossman & Jian-Kui Liu & Wen-Jing Li &
Don-Qing Dai & Jayarama D. Bhat & E. B. Gareth Jones & Eric H. C. McKenzie &
Jian-Chu Xu & Kevin D. Hyde
#
School of Science 2014
Abstract T ubeufiaceae is based on the generic type T ubeufia,
which is characterized by superficial, oval and bright ascomata,
bitunicate asci, mostly long fusiform to filiform, transeptate
ascospores and hyphomycetous asexual states with
helicosporous conidia. Most species in this family are saprobic
on terrestrial woody substrates and some are aquatic. Their
distinct morphology as well as combined LSU, SSU and
TEF1 sequence analysis show that T ubeufiaceae should be
accommodated in a new order Tubeufiales, which is introduced
in this paper. Phylogenetic analyses of combined LSU and ITS
sequences were used to resolve genera and species within the
family T ubeufiaceae. In this study, we examine and incorporate
sexual and asexual states of genera in T ubeufiales to provide a
modern treatment, based on single names. An epitype for
Tubeufia javanica, the type species of Tubeufia, is designated
and represents Tubeufia sensu stricto. The genera
Acanthophiobolus, Acanthostigma, Boerlagiomyces,
Chlamydotubeufia, Kamalomyces, Podonectria, Thaxteriella
and Thaxteriellopsis are accepted, Acanthostigmina is reinstated ,
and the asexual genera Aquaphila, Helicoma, Helicomyces,
Helicosporium and Tamhinispora are accepted in T u beufiaceae.
Three new genera Acanthohelicospora, Helicangiospora and
Neoacanthostigma are introduced. The genus Bifrontia is added
to the family based on morphological similarity. The incongru-
ous morphological genera Acanthostigmella, Amphinectria,
Chaetocrea, Chaetosphaerulina, Glaxoa, Malacaria,
Melioliphila, Paranectriella, Puttemansia, Rebentischia and
Ur edin ophila are excluded from T ubeufiaceae despite having
characteristic ascomata with setae and multiseptate long spores.
A key to genera accepted in Tubeufiaceae is provided.
Keywords Dothideomycetes
.
Helicospores
.
New genus
.
Phylogeny
.
Taxonomy
.
Tubeufiaceae
Electronic supplementary material The online version of this article
(doi:10.1007/s13225-014-0304-7) contains supplementary material,
which is available to authorized users.
S. Boonmee
:
W.<J. Li
:
D.<Q. Dai
:
J.<C. Xu
:
K. D. Hyde (*)
Key Laboratory for Plant Diversity and Biogeography of East Asia,
Kunming Institute of Botany, Chinese Academy of Science,
Kunming 650201, Yunnan, China
e-mail: kdhyde3@gmail.com
S. Boonmee
:
W.<J. Li
:
D.<Q. Dai
:
J.<C. Xu
:
K. D. Hyde
World Agro forestry Centre East and Central Asia Office, 132 Lanhei
Road, Kunming 650201, China
S. Boonmee
:
J.<K. Liu
:
W.<J. Li
:
D.<Q. Dai
:
K. D. Hyde
Institute of Excellence in Fungal Research, Mae Fah Luang
University, Muang, Chiang Rai 57100, Thailand
S. Boonmee
:
J.<K. Liu
:
W.<J. Li
:
D.<Q. Dai
:
K. D. Hyde
School of Science, Mae Fah Luang University, Muang, Chiang
Rai 57100, Thailand
S. Boonmee
:
A. Y. Rossman
Systematic Mycology and Microbiology Laboratory, USDA-ARS,
Beltsville, MD 20705, USA
J. D. Bhat
Formerly at Department of Botany, Goa University, Goa 403 206,
India
E. B. G. Jones
Department of Botany and Microbiology, King Saudi University,
Riyadh, Saudi Arabia
E. H. C. McKenzie
Landcare Research, Auckland, New Zealand
DOI 10.1007/s13225-014-0304-7
Received: 13 July 2014 /Accepted: 25 August 2014 /Published online: 26 September 2014
Fungal Diversity (2014) 68:239–298
Introduction
The family Tubeufiaceae was established in Pleosporales
by Barr (1979) based on the generic type Tubeufia and
included five genera i.e. Letendraeopsis, Melioliphila,
Podonectria, Rebentischia and Thaxteriella,whileare-
cent treatment by Boonmee et al. (2011)included20
genera (Table 1). Allonecte, Byssocallis, Letendraeopsis
and Taphrophila were considered atypical and excluded
from the family. Most genera of Tubeufiaceae have
uniloculate, superficial, pigmented ascomata e.g. pale
brown, brown, and dark brown to black, multi-celled,
hyaline ascospores, and produce helicosporous asexual
states (Tsui et al. 2006; Boonmee et al. 2011). For a
comprehensive review of Tubeufiaceae, see Boonmee
et al. (2011).
Molecular techniques are being used to establish taxonom-
ic relationships among genera in Dothideomycetes (Lutzoni
et al. 2004; Hibbett et al. 2007; H yde and Zhang 2008;
Lücking 2008; Brock et al. 2009; Eberhardt 2010;
Figueiredo et al. 2010; Liu et al. 2011, 2012; Zhang et al.
2012) and establish their higher level placement in a natural
classification system (Hyde et al. 2013; Ariyawansa et al.
2014a, b; Hongsanan et al. 2014; Phookamsak et al. 2014;
Thambugala et al. 2014; Wijayawardene et al. 2014). Some
studies on Tubeufiaceae have also incorporated molecular
analyses and these results were reviewed in Boonmee et al.
(2011). Basically, in recent phylogenetic a nalyses of
T ubeufiaceae, species in different sexual and asexual genera
of T ubeufiaceae were scattered throughout the trees (Tsui and
Berbee
2006;Tsuietal.2006). Therefore generic concepts
were difficult to resolve and it proved impossible to establish
which asexual genera were linked to sexual morphs. Morpho-
logical characterization and phylogenetic reconstruction have
shown that members of T ubeufiaceae form a monophyletic
clade in Dothideomycetes, and are a sister lineage to
Pleosporales (Tsui and Berbee 2006; Schoch et al. 2006,
2009; Boonmee et al. 2011; Zhang et al. 2011, 2012; Hyde
et al. 2013). The objective of the present paper is to introduce a
new order Tubeufiales to accommodate the monophyletic
family T ubeufiaceae, which has previously been classified in
Pleosporales. Integration of sexual and asexual names in
T ubeufiaceae is now possible and each genus should have a
single name (Taylor 2011;Gamsetal.2012; McNeill et al.
2012; Hawksworth et al. 2013). In this paper we epitypify or
designate authentic sequenced specimens for type species of
genera in order to stabilize the understanding of each genus. In
this way we are able to connect asexual and sexual genera
Tabl e 1 Various treatments of Tubeufiaceae, Tubeufiales. A question mark (?) indicates that the position of the taxon is uncertain
Barr 1979 Barr 1980 Rossman 1987 Kirk et al. 2001 Lumbsch and Huhndorf 2010 Boonmee et al. 2011 In this study
Letendraea Allonectria Allonectria Acanthophiobolus Acanthostigma Acanthostigma Acanthohelicospora
Melioliphila Boerlagiomyces Boerlagiomyces Acanthostigmella Acanthophiobolus Acanthophiobolus Acanthophiobolus
Podonectria Byssocallis Byssocallis Allonecte Acanthostigmella ?Acanthostigmella Acanthostigma
Rebentischia Letendraea Letendraea Amphinectria Allonecte ?Amphinectria Acanthostigmina
Thaxteriella Melioliphila Malacaria Boerlagiomyces ?Amphinectria Aquaphila Aquaphila
Tubeufia Paranectriella Melioliphila Borinquenia Boerlagiomyces ?Boerlagiomyces Bifrontia
Podonectria Paranectriella Byssocallis Byssocallis ?Chaetocrea ?Boerlagiomyces
Puttemansia Podonectria Chaetocrea Chaetocrea Chaetosphaerulina Chlamydotubeufia
Rebentischia Puttemansia Glaxoa Chaetosphaerulina ?Glaxoa Helicangiospora
Tubeufia Rebentischia Letendraea Glaxoa Kamalomyces Helicoma
Tubeufia Letendraeopsis Letendraeopsis ?Malacaria Helicomyces
Uredinophila Malacaria Malacaria ?Melioliphila Helicosporium
Melioliphila Melioliphila ?Paranectriella Kamalomyces
Paranectriella Paranectriella Podonectria Neoacanthostigma
Podonectria Podonectria ?Puttemansia Podonectria
Puttemansia Puttemansia Rebentischia Tamhinispora
Rebentischia Rebentischia Thaxteriella Thaxteriella
Taphrophila Taphrophila Thaxteriellopsis Thaxteriellopsis
Thaxterina Thaxteriella Tubeufia Tubeufia
Tubeufia ?Thaxteriellopsis ?Uredinophila
Uredinophila Thaxterina
Tubeufia
Uredinophila
240
Fungal Diversity (2014) 68:239–298
through molecular analysis and determine priorities for the
linked genera.
Materials and methods
Examination of herbarium specimens
The generic types or authentic specimens of tubeufiaceous
genera, namely Acanthostigmella, Acanthostigmina,
Bifrontia, Chaetocrea, Helicoma, Helicomyces,
Helicosporium, Malacaria, Paranectriella, Puttemansia,
Rebentischia, Thaxteriella, Tubeufia and Uredinophila were
obtained from the repositories of U. S. National Fungus Col-
lections (BPI), Farlow Reference Library and Herbarium of
Cryptogamic Botany in Harvard University (FH), Kew Royal
Botanic Gardens (K), Natural History Museum, University of
Oslo, Botanical Museum (O) and Naturhistorisches Museum
Wien (W). Morphological features were examined and pho-
tomicrographs made using Carl Zeiss Microscopes, measure-
ment confirmed using software of Micro Imaging GmbH.
AxioVs40 V 4.8.2.0 (2006–2010).
Fresh collections
Decaying wood samples were randomly collected from sam-
pling sites in the forests of Chiang Mai a nd Chiang Rai
provinces, northern Thailand. Samples were taken to the
laboratory in Zip-lock plastic bags. The material was exam-
ined under a Motic SMZ 168 series microscope. Micro-
morphological structures were photographed using a Nikon
ECLIPSE 80i compound microscope fitted with a Canon
450D digital camera and measurements made using Tarosoft
(R) Image Frame Work program. Figures were processed with
an Adobe Photoshop CS3 Extended version 10.0 (Adobe
Systems Inc., USA). Type materials are deposited in the
herbarium of Mae Fah Luang University (Herb. MFLU ),
Chiang Rai, Thailand, with isotypes in th e New Zealand
Fungal and Plant Disease Collection (PDD) and cultures in
Mae Fah Luang University Culture Collection (MFLUCC),
BIOTEC Culture Collection (BCC), Thailand and Internation-
al Collection of Microorganisms from Plants (ICMP), New
Zealand; some isolates are maintained in IFRD culture collec-
tion, International Fungal Research & Development Centre,
Kunming, China, under the Material Transfer Agreement No.
4/2010 (MTA).
Isolation of fungi
Single spore isolates were made on water agar (WA) or malt
extract agar (MEA, Difco Laboratories, Detroit, Michigan,
USA) and germinating spores were aseptically transferred to
fresh MEA plates and incubated at 28 C for 7 days following
the methods of Boonmee et al. (2011) and Chomnunti et al.
(2011, 2014). Cultures were grown for 1–2monthsandmor-
phological characters such as colour, colony shape, and tex-
ture recorded. The cultures were checked for asexual states
after 30–60 days of growth.
DNA sequences
Genomic DNA was extracted from fungal mycelium grown
on MEA at 28 C for 30 days. Four genes were amplified with
universal primers, namely the internal transcribed spacer (ITS:
ITS5/ITS4) region of ribosomal DNA, large subunit nuclear
ribosomal DNA (LSU: LROR/LR5), small subunit nuclear
ribosomal DNA (SSU: NS1/NS4) and the translation elonga-
tion factor-1 alpha (TEF1a: TEF1 983/2218R) (White et al.
1990; Vilgalys and Hester 1990; Schoch et al. 2009). The
PCR products were purified and sequenced with the same
primers.
Molecular data analyses
Phylogenetic trees
BLAST search of new sequences were performed to verify the
identities of species with sexual and asexual states,
Tubeufiaceae in the GenBank database (Kodsueb et al.
2004;TsuiandBerbee2006;Tsuietal.2006;Promputtha
and Miller 2010; Boonmee et al. 2011; Sánchez et al. 2012).
Details of sequences are provided in Supplementary Table 1.
Two datasets were carried out for phylogenetic analyses of
tubeufiaceous fungi. The first dataset aims to deal with the
classification of Tubeufiales wihin Dothideomycetes
(Chomnunti et al. 2012; Liu et al. 2012;Ariyawansaetal.
2013;Hydeetal.2013
; Pérez-Ortega et al. 2014), which used
multi-gene LSU, SSU and TEF, comprising 141 taxa includ-
ing the outgroup (Schismatomma dec olorans (Turner &
Borrer ex Sm.) Clauzade & Vězda DUKE 0047570,
Arthoniomycetes). The second dataset focus on phylogenetic
relationships within taxa of the Tubeufiaceae (Tsui et al. 2006;
Promputtha and Miller 2010; Boonmee et al. 2011;Sánchez
et al. 2012), focued by using the combination of LSU and ITS
sequences, consisting of 68 taxa including the outgroup
(Botryosphaeria dothidea (Moug.) Ces. & De Not.
CBS115476, Botryosphaeriales). Sequences and alignments
were prepared using BioEdit (Hall 1999). Alignment data
were parsed with Gblocks (Castersana 2000) following the
default setup. Phylogeny website tools “ALTER” (Glez-Peña
et al. 2010) was used to transfer the alignment file for RAxML
analysis. The reconstruction of the maximum likelihood (ML)
analysis usingRAxML v. 7.2.8 (Stamatakis et al. 2008)aspart
of the “RAxML-HPC BlackBox on TG tool” performed at the
241
Fungal Diversity (2014) 68:239–298
CIPRES Science Gateway v. 3.3 ( http://www.phylo.org/
portal2/;Milleretal.2010). All free model parameters will
be estimated by RAxML and ML estimate of 25 per site rate
categories. Final ML search were conducted under the GTRG
AMMA model.
The Bayesian command was generated using FaBox 1.41
(Villesen 2007). To determine Bayesian posterior probability
was performed using Old MrBayes 3.1.2 on XSEDE, param-
eters setting of 2 parallel runs, 4 chains, carried out for 4 000
000 generations, sample frequency every 1,000 generations,
and all other parameters were left as default (Huelsenbeck and
Ronquist 2001). The 50 % majority rule consensus tree was
constructed from the remaining trees in Treeview (Page 1996).
Results and discussion
Phylogenetic study
The phylogenetic tree was built from a multigene analyses of
140taxainofDothideomycetes with Schismatomma
decolorans (Arthoniomycetes) as the outgroup (Fig. 1). The
datas et comprises 141 taxa, with 3,415 characte rs, 1,682
constant characters, 1,368 parsimony informative characters
and 365 variable characters which are parsimony uninforma-
tive (TL=8531, CI=0.317, RI=0.646, RC=205, HI=683).
Twenty orders are recognized in Dothideomycetes, and a
distinct calde was formed, namely Tubeufiales, and has a close
relationship with the order Patellariales and
Botryosphaeriales, which shows the same phylogenetic rela-
tionships with the previous studies of Schoch et al. (2009)and
Hyde et al. (2013). However, it is far removed from
Pleosporales where the family Tubeufiaceae was previously
assigned (Barr 1979). The recently introduced order
Abrothallales (Perez-Ortega et al. 2014) was included in our
analysis. Perez-Ortega et al. (2014)statedthatAbr othallales
formed a sister group of Jahnulales and close with
Patellariales. In our phylogenetic analysis, Abrothallales
formed a sister group with Lichenoconiales and is close to
Jahnulales. However, it is not related to Patellariales which is
always close with T ubeufiaceae.
The phylogenetic tree in Fig. 2 resulted from the molecular
analysis of LSU and ITS combined data. The data setup
consists of 67 taxa, with 1,096 characters, 777 characters are
constant, 244 were parsimony informative and 75 variable
characters were parsimony uninformative (TL=1091, CI=
0.420, RI=0.731, RC=0.307, HI=0.580). The taxa in the tree
can be divided into 13 clades (namely A-M), each receiving
moderate to strong statistical support and having representa-
tive sharing distinguished morphology (Fig. 2). Phylogenetic
trees were drawed in Treeview (Page 1996) and MEGA 5
(Tamura et al. 2011).
Clade A is represented by taxa of Helicoma sensu stricto
(86 % BS and 1.00 PP), which includes an authentic strain of
the type species of Helicoma (H. muelleri Corda). Putatively
named strains of Helicosporium linderi R.T. Moore (NBRC
9207), T. paludosa (P. Crouan & H. Crouan) Rossman
(HKUCC 9118, ANM 196, ANM 1169) and Thaxteriella
helicoma (W. Phillips & Plowr.) J.L. Crane, Shearer & M.E.
Barr (JCM 2739, UBCF 13877) clustered in this clade and we
recognize them in Helicoma in the phylogenetic tree (Fig. 2),
but do not formally synonymize them as we have not seen
voucher material. Helicoma rugosa asexual morphs formed in
cultures of Thaxteriella helicoma (JCM 2739 and UBCF
13877). Tubeufia khunkornensis Boonmee & K.D. Hyde
(MFLUCC 10–0119, ex-type strain), T. inthanonensis
Boonmee & K.D. Hyde (MFLUCC 11–0003, ex-type strain)
and T. miscanthi W.H. Hsieh, Chi Y. Chen & Sivan.,
epitypified here are synonymized under Helicoma. Helicoma
chiangraiense, H. fagacearum and H. siamense formed dis-
tinct groups at the base of Clade A, and we therefore introduce
them as new species of Helicoma.
Clade B is the genus Thaxteriellopsis sensu stricto repre-
sented by four stains of the type species T. lignicola Sivan.,
Panwar & S.J. Kaur, of which MFLUCC 10–0124 is the ex-
epitype.
Clade C represents the monotypic genus Helicangiospora,
which is introduced in this study. Its morphology is similar
with Acanthostigma, but it is phylogenetically distinct (Clade
E). In addition, its asexual morph is unique in producing
conidial helicospores borne in a capsule, which distinguishes
this genus from all known genera of T ubeufiaceae.
Clade D is Helicosporium sensu stricto (84 % BS and 1.00
PP) represented by helicosporium-like taxa. The type species
Helicosporium vegetum Nees is represented by four strains
(NBRC 30345, CBS 941. 72, BCC 3332 and BCC 8125),
however, these four strains did not cluster together. Strain
CBS 941. 72 has been considered to be the authentic strain
of H. vegetum, and at least two strains (BCC 3332 and BCC
8125) are probably not H. vegetum sensu stricto. Two strains
of Tubeufia cerea (Berk. & M.A. Curtis) Höhn. (CBS 254.75
and NBRC 9014) also cluster with H. vegetum sensu stricto
(99 % BS and 1.00 PP) and are thus synonymised.
Helicosporium guianense Linder (CBS 269.52) also clusters
with H. vegetum sensu stricto
and is probably wrongly named.
An ex-type strain of Acanthostigma patagonicum R.M.
Sánchez, A.N. Mill. & Bianchin. (MVB 573 BBB) and
Helicoma vaccinii Carris (CBS 216.90) clustered in this clade
Fig. 1 RAxML phylogenetic placement of the new order Tubeufiales
and allied orders within the Dothideomycetes. RAxML bootstrap support
values≥50 % (BS) and Bayesian posterior probabilities≥ 0.95 (PP) are
shown at the nodes (values below these thresholds not shown). The tree
was rooted with Schis matomma decolorans DUKE 0047570
(Arthoniomycetes). Ex-type strains are in bold
242
Fungal Diversity (2014) 68:239–298
243
Fungal Diversity (2014) 68:239–298
Fig. 1 (Continued)
244
Fungal Diversity (2014) 68:239–298
and we therefore rename them as Helicosporium in the phy-
logenetic tree (Fig. 2). We do not formally synonymize them
as the basal grouping of these two species in this Clade is
variable and the morphology differs.
Clade E represents Acanthostigma sensu stricto with three
taxa, including the type species A. perpusillum De Not. This
clade always clustered together with Helicosporium sensu
stricto, and the position is stable in indivual gene analysis
(data not shown).
Clade F with seven taxa represents Tubeufia sensu stricto
which is the type genus of T ubeufiaceae. The type species,
T. javanica Penz. & Sacc. is epitypified and a new species
T. chiangmaiensis is introduced in this study. With the excep-
tion of T. cylindrothecia (Seaver) Höhn. (BCC 3559), the
other four taxa from the study of Tsui et al. (2006) were
identified as species of Helicomyces and Helicoma.Inour
phylogenetic analysis, these strains clustered together and
formed a distinct c lade, therefore, we rename them as
Tubeufia species in the phylogenetic tree (Fig. 2), but do not
formally synonymize them as we have not seen voucher
material. The morphology of Tubeufia sensu stricto are de-
tailed based on T. javanica and T. chiangmaiensis.
Clade G (Helicomyces sensu stricto) is phylogenetically
close to the Tubeufia sensu stricto clade and is represented by
an authentic strain of the type species, H. roseus Link (CBS
283.51), plus two cultures of Tubeufia paludosa (CBS
120503, ep itypified he rein). Tubeufia pa ludosa is also
synonymised under Helicomyces. The putativel y name d
strains of Helicosporiu m indicum P.Rag. Rao & D. Rao
(CBS 374.93) and H. talbotii Goo s (MUCL 33010) also
cluster with Helicomyces sensu stricto. These are renamed
as Helicomyces species in the phylogenetic tree (Fig. 2), but
they are not formally synonymized as we have not seen
voucher material.
Clade H represents the monotypic genus Tamhinispora,
which was introduced by Rajeshkumar and Sharma (2013).
Its morphology is similar with Chlamydotubeufia, but differs
by its ovoid, dictyoseptate conidia with apical appendages.
This genus is the second group of Tubeufiales known only
from freshwater, and the other one is Aquaphila. They differ
as Aquaphila produces sickle-shaped conidia, while
Tamhinispora has ovoid, dictyoseptate conidia with apical
appendages. The sexual morph of Tamhinispora is unknown.
Clade I represents the genus Chlamydotubeufia incuding
three species, which form a well-supported clade (85 % BS
and 1.00 PP). Both asexual and sexual morphs occur in this
genus, and it is a well-defined genus. Chlamydotubeufia
chlamydosporum (Shearer) Boonmee & K.D. Hyde and
C. huaikangplaensis Boonmee & K.D. Hyde cluster with
100 % BS and 1.00 PP but we do not synonymise them here
as the tree is not well-populated and it is not clear that are
morphologically identical (Shearer 1987; Boonmee et al.
2011).
Clade J represents the freshwater genus Aquaphila sensu
stricto. The type species A. albicans Goh, K.D. Hyde & W.H.
Ho and its tubeufia-like sexual state (Tubeufia asiana Sivichai
& K.M. Tsui) formed a well-supported clade and has a close
relationship with Chlamydotubeufia. Based on our phyloge-
netic analysis, we synonymize Tubeufia asiana unde r
Aquaphila albicans.
Clade K represents the new genus Acanthohelicospora
with A. pinicola as the type species. Three species with two
strains from Acanthostigma and two species from
Helicosporium are renamed Acanthohelicospora in Fig. 2
based on both morphological and phylogenetic studies. How-
ever, as these are not ex-type strains they are not formally
transferred. This genus produces a unique helicosporous asex-
ual state that distinguishes it from other genera in
T ubeufiaceae.
Clade L represents Acanthostig mina sensu stricto with
three strains of the type species A. minuta (Fuckel) Clem. &
Shear and two strains of Acanthostigma multiseptatum Promp.
& A.N. Mill. There are no ex-type strains A. minuta.We
synonymize Acanthostigma multiseptatum (one ex-type
strain) under Acanthostigmina based on the phylogenetic
result.
Clade M is basal in Tubeufiaceae and represents the new
genus Neoacanthostigma with N. fusiforme as the type spe-
cies. All the three species are represented by ex-type strains.
The species N. septoconstrictum and N. filiforme were previ-
ously named Acanthostigma, however, their morphological
characters, such as the asci and ascospores, differ from these
genera. Thus they are synonymized under Neoacanthostigma
here.
Taxonomy
Tubeufiales S. Boonmee & K.D. Hyde,
ord. nov., Index
Fungorum number: IF 550704, Facesoffunginumber: FoF
00203
Saprobic, common on decorticated or decaying woody and
herbaceous substrates, often associated with decaying fungi,
less common on leaves, terrestrial and aquatic habitats, wide-
spread in temperate to trop ical regio ns. Sexual state:
Ascomata completely superficial, seated on a subiculum, uni-
locular, globose-subglobose or clavate to obovate, soft-tex-
tured, solitary to gregarious, partially grouped, translucent or
not, pale brown, brown to black, minutely papillate and with
ostiolar, collapsing cupulate, laterally or not when dry, with
radiating mycelium or appendages at base, with or without
setose or hairy appendages. Peridium somewhat thickened,
mostly composed of cells of textura angularis,thick-walled
cells, pale yellow, brown, dark brown to black externally, with
thin layers of textura prismatica inwardly, cells narrow, slight-
ly elongate, hyaline, pale brown to brown. Hamathecium
comprising numerous filiform, septate, branched, sometimes
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Fungal Diversity (2014) 68:239–298
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Fungal Diversity (2014) 68:239–298
anastomosing, hyaline pseudoparaphyses, embedded in a ge-
latinous matrix. Asci 8-spored, bitunicate, fissitunicate, sac-
cate or cylindrical-clavate, sometimes broadly oblong-
subclavate, with or without an apically rounded, distinct ocu-
lar chamber, with or without a distinct pedicel. Ascospores 2–
3-seriate to fasciculate in ascus, elongate, cylindric-
subfusiform to narrowly oblong, tapering towards narrow,
subacute ends, distinctly multiseptate, hyaline to pale brown,
smooth or minutely verrucose. Asexual states: hyphomy-
cetous; helicosporous, chlamydosporous and
phragmosporous. Conidiophores mononematous, macro- to
micronematous, erect, flexuous, septate, pale brown, brown
to dark brown. Conidiogenous cells holoblastic, mono- or
polyblastic, integrated or discrete, terminal or intercalary.
Conidia usually elongate, filiform to fusiform, curved, heli-
coid with varying number of coils, septate, sometimes
dictyosporous, phragmosporo us, hyaline to variously
coloured, smooth-walled to verrucose.
Ordinal type: Tubeufiaceae M.E. Barr, Mycologia 71: 948
(1979), Facesoffunginumber: FoF 00204
Monotypic, characters same as for Tubeufiales.
Family type: Tubeufia Penz. & Sacc.
Notes: Below we list the genera accepted in the
T ubeufiaceae. Acceptance is based on morphological charac-
teristics and/ or molecular data. Generic names based on
sexual or asexual types are treated equally.
Key to genera of Tubeufiaceae
1.Ascomatapresent ........................... 0.2
1.Ascomatalacking .......................... 0.15
2. Ascomata with setose appendages or flexuous hyphae,
seateddirectlyonsubstrate .................... 0.3
2. Ascomata without appendages, seated on a basal
subiculum ......................0.12
3.Asexualstateknowninculture ................. 0.4
3.Asexualstatenotknown ...................... 0.7
4. Conidia dictyochlamydosporous, black . .Chlamydotubeufia
4.Conidiahelicosporous,hyalinetopalebrown ...... 0.5
5. Conidia born in sheath, helicoma-like . . . . Helicangiospora
5.Conidiabornonhyphae,helicosporium-like ....... 0.6
6. Conidiophores mononematous . . ....Acanthohelic ospora
6. Conidiophores micronematous . . . . . Neoacanthostigma
7.Ascosporesfiliform,spiralinasci .....Acanthophiobolus
7. Ascospores fusiform, clavate-fusiform, cylindric-fusoid, 2–
3-seriateinasci ............................. 0.8
8. Ascospores muriform, dictyoseptate . . . Boerlagiomyces
8. Ascospores phragmosporous, trans-septate . . . . . . . . 0.9
9. Ascospores greater than 30-septate . . . . . . Kamalomyces
9.Ascosporeslessthan30-septate................ 0.10
10. Ascospores equally 5-septate . . . . . . . . Thaxteriellopsis
10.Ascosporesgreaterthan5-septate ..............0.11
11. Ascomata greater than 200μm diam; ascospores hyaline
to pale brown ..Acanthostigmina
11. Ascomata less than 200μ m diam; as cospo res hya-
line ...................Acanthostigma
12.Ascomataonscaleinsects ............. Podonectria
12. Ascomata on decaying or rotting wood . . . . . . . .0.1312
13. Ascospore s allantoid or vermiform, equa lly 7-
septate ..................Thaxteriella
13. Ascospores elongated cylindrical-subfusiform, greater
than7-septate ...........................0.1413
14. Ascomata pale yellow to pale brown; with numerous
pseudoparaphyses..................... Tubeufia
14. Ascomata dark brown to black; lacking
pseudoparaphyses .............. Bifrontia
15. Con idia dictyosporous, apically appendaged, heavily
pigmented .......................Tamhinispora
15. Conidia phragmosporous, helicosporous, light
pigmented .....................0.16
16. Conidia fusoid to sickle-shaped, slightly curved, always
aquatic .............................Aquaphila
16.Conidiacoiled,aquaticorterrestrial ........... 0.17
17. Conidiophores absent; conidia borne directly on
hyphae . . . ............... Helicomyces
17. Conidiophores present, dark pigmented . . . . . . . . . 0.18
18. Conidia coiled 1–1½ times; conidial filament greater than
6μm wide ...........................Helicoma
18. Conidia coiled 3½–4½ times; conidial filament less than
6μm wide.......................Helicosporium
Accepted genera
Tubeufia Penz. & Sacc., Malpighia 11(11–12): 517 (1898),
Facesoffunginumber: FoF 00063
Saprobic on decorticated or decaying woody and herba-
ceous substrates, often associated with other fungi, in terres-
trial habitats, widespread in temperate to tropical regions.
Sexual state: Ascomata superficial, seated on a subiculum,
spherical to clavate-obovate, oval to ellipsoid, bright to
yellowish-brown when young, brown to dark brown when
mature or dry, apex rounded, narrow at base, darkly
pigmented, occasionally collapsing when dry. Peridium com-
prising pale brown cells of textura angularis,andsmall
textura prismatica subhyaline cells at innermost layers.
Hamathecium comprising numerous filiform, septate,
branched, hyaline pseudoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, saccate or cylindric-clavate, with or
without an ocular chamber. Ascospores fasciculate in ascus,
elongate cylindric-subfusiform or narrowly oblong, greater
Fig. 2 Phylogenetic representation of Tubeufiaceae and its members.
ThetreeisrootedwithBotryosphaeria dothidea CBS115476
(Botryosphaeriales). RAxML bootstrap support values≥70 % (BS) and
Bayesian posterior probabilities ≥ 0.95 (PP) are shown at the nodes and
bold (values below these thresholds not shown). Ex-type strains are in
bold. The type species of each genus is in blue. Thick lines on branches
determine clades that are resolved in the RAxML consensus
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Fungal Diversity (2014) 68:239–298
than 5-septate, not constricted at septa, hyaline to pale brown,
smooth-walled. Asexual state: hyphomycetous,
helicosporous.
Type species: Tubeufia javanica Penz. & Sacc.
Notes: Tubeufia sensu stricto (Clade F, Fig. 2)comprises
species related to T. javanica having light-coloured, superfi-
cial, smooth-walled ascomata and hyaline to pale brown,
elongate cylindric-subfusiform or narrowly oblong asco-
spores. In this paper, we describe, illustrate and designate an
epitype specimen for T. javanica (MFLUCC 12–0545), and
introduce T. chiangmaiensis as a new species. By designating
an epitype with sequence data we aim to stabilize the genus so
that related species and asexual states can be placed in this
genus. The asexual morphs of Tubeufia sensu stricto are
helicosporous as four species with helicomyces-like and
helicoma-like conidia cluster in Clade G, i.e. Tubeufia
cylindr othecia, T. intermedium, T. lilliputeus, T. roseus and
T. talbotii. However, we could not examine the characters of
these strains deposited in GenBank and t hus do not
formally synonymize them. With the introduction of
one new species below, Tubeufia sensu stricto presently
comprises only two species and severa l putative ly
named strains in GenBank.
Tubeufia javanica Penz. & Sacc., Malpighia 11(11–12):
517 (1898), Facesoffunginumber: FoF 00063 Fig. 3
Holotype: INDONESIA, Java, Tjibodas, on withered culm
of Bambusa emar cidis,2Mar1987(PAD)
Epitype designated here:MFLU13–0371
Saprobic on culms or sheaths of bamboo. Sexual state:
Ascomata 376–505(−550) μm high×(200-)224–260(−280)
μm diam., superficial, seated on a subiculum, solitary, some-
times gregarious, oblong, subclavate to obclavate, cream-
white to yellowish when young, pale brown to brown when
mature, apex rounded, base narrow, brown to dark brown,
collapsing when dry, compressed subiculum hyphae, 2.5–
5μm wide, partially branched, septate, dark brown, and irreg-
ular. Peridium 15–25μm wide, comprising light brown cells
of
textura angularis, and small textura prismatica subhyaline
cells inwardly. Hamathecium comprising numerous, 1–2μm
wide, filiform, septate, branched, hyaline pseudoparaphyses.
Asci (140-)164– 221(− 230)×14– 18(− 22) μ m
(
x¼ 194 16μm , n=20), 8-spored, bitunicate, fissitunicate,
cylindrical, short pedicellate, apically rounded, with an ocular
chamber. Ascospores ( 95-)100.5–117(−120)×4– 5 μm
(
x¼ 109 4:5μm , n=20), fasciculate, broad filiform, cylin-
drical to long subfusiform, elongate, ends rounded, 14–17-
septate, not constricted at septa, hyaline to pale yellow or
brown, smooth-walled. Asexual state: Unknown.
Cultural characteristics: Ascospores germinating on PDA
within 24 h and germ tubes produced from both ends. Colonies
growingslowlyonPDA,reaching4.5mmin2weeksat28C,
effuse, velvety to hairy, edge fimbriate, olive to olive brown,
dark brown in PDA media. Mycelium superficial and partially
immersed, branched, septate, pale brown to olivaceous brown,
smooth, asexual spores not formed within 60 days.
Material examined: THAILAND, Chiang Rai, Muang,
Khun Korn Waterfall, on dead culm of bamboo Kunth ex
Dumort (Poaceae), 31 July 2012, Dong-Qin Dai,
DDQ00239 (MFLU 13–0371, epitype of Tubeufia javanica
designated here; PDD 104450, iso-epitype), ex-type living
culture = MFLUCC 12–0545 = ICMP 20067; Chiang Rai,
Muang, Khun Korn Waterfall, on dead culm-sheath of bam-
boo, 21 July 2013, S. Boonmee KK-11 (MFLU 14–0222),
living culture = MFLUCC 14–0438.
Notes: We requested the type speciman from PAD and BO
but this is not present in either herbarium and therefore must
be presumed lost. We therefore designate our fresh collection
as epitype to stabilize applications of the genus and species.
The epitype is representative of the iconotype and description
provided in the protologue, and the location in Asia and
bamboo host are also appropriate for epitypification (Penzig
and Saccardo 1904; Boonmee et al. 2011).
Tubeufia chiangmaiensis S. Boonmee & K.D. Hyde, sp.
nov. Index Fungorum number: IF 550705,
Facesoffunginumber: FoF 00172 Fig. 4
Etymology: named after Chiang Mai, the location where it
was collected.
Holotype:MFLU11–1149
Saprobic on dead wood . Sexual state: Ascomata
(200-)238– 324 μ m high × (180-) 227– 269 μ m diam.
(
x¼ 266 228μm ), superficial, solitary, scattered, globose-
subglobose to ovate, dark brown, with hyphae developing
from ascomatal base onto substrate, slightly flat at the apex,
browntodarkbrown,ostiolate,collapsingwhendry.
Peridium 25–29μm wide, comprising 4–5layersofbrown
to reddish-brown cells of textura angularis, and small,
subhyaline to light brown cells of textura prismatica inwardly.
Hamathecium comprising numerous, ca. 1–1.5μm wide, fili-
form, septate, branched, hyaline pseudoparaphyses. Asci
(114.5-)120–137×11–14(−16) μm (
x¼ 127 13μm , n=20),
8-spored, bitunica te, fissitu nicat e, elongate cylin dri cal to
slightly c lavate, apedicell ate, thick-walled, rounded at
apex, with an ocular chamber, tapering towards narrow
base, sessile. Ascospores 46–53(−55.5)×(3.5-)4–4.5μm
(
x¼ 50 4μm , n=20), overlapping 2–3-seriate, cylindric-
fusiform, tapering toward ends, with pad-like mucilage at both
ends, straight to slightly curved, 7-septate, not constricted at
Fig. 3 Tubeufia javanica (MFLU 13–0371, epitype). a, b Ascomata
formed on natural substrate. c Close up of pale reddish-brown ostiole. d
Side view of ascoma located on a subiculum. e L.S. of ascoma. f
Peridium. g Pseudoparaphyses. h–j Asci. k Ruptered ascus. l–o Asco-
spores. p Germinating ascospore. q Dark brown colonies on PDA. r, s
Growth on plant substrates and aerial hyphae in culture. Scale bars: a =
500μm, b–c =200μm, d–e, h–k =100μm, f =40μm, e =5μm, l–p =50,
q–r =10mm,s =20μm
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septum, hyaline, pale brown when mature, smooth-walled.
Asexual state:Unknown.
Cultural characteristics: Ascospores germinating on MEA
within 12 h and ge rm tubes produced from both ends.
Colonies growing on MEA, reaching 5 mm in 7 days at 28
C, mycelium partly superficial, partly immersed, slightly ef-
fuse, radially striate, with fimbriate edge, dark-coloured; asex-
ual spores not formed within 60 days.
Material examined: THAILAND, Chiang Mai, Mae
Taeng, Mushroom Research Center, N19°17.123′ E
98°44.009′,elev.ca. 900 msl., on dead wood of an unidenti-
fied tree , 23 June 2011, Saranyaphat Boonmee MRC-06
(MFLU 11–1149, holotype); ex-type living cultures =
MFLUCC 11–0514 = BCC 52386 = ICMP 20074.
Notes: Tubeufia chiangmaiensis has morphological char-
acters typical of the genus, such as brown, globose-
Fig. 4 Tubeufia chiangmaiensis (MFLU 11–1149, holotype). a
Ascomata on substrate. b L.S. of ascoma. c Peridium. d Hamathecium
and pseudoparaphyses. e–g Asci. h–j Ascospores with mucilaginous pad-
like appendage at both ends, marked by arrows. k Germinating
ascospore. l, m Colonies on MEA from surface and reverse. Note dark
brown colonies. Scale bars: a–b =100μm, c, h–k = 30 mm, d =
5μ m, e–g =50μm, l–m =10mm
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subglobose ascomata with basal mycelium, which darken
when dry, fissitunicate, elongated, cylindrical-clavate asci,
and cylindrical-fusiform, 7-transeptate, hyaline to pale brown
ascospores, with pad-like mucilage at both ends. Tubeufia
cylindrothecia has oblong translucent peridial ascomata, as-
cospores lack mucilaginous pad-like appendages and are
smaller when compared to Tubeufia chiangmaiensis (Seaver
1909). Tubeufia chiangmaiensis differs from T. javanica in
ascomatal and ascospore features (Penzig and Saccardo 1904;
Rossman 1977; Sivanesan 1984, Fig. 3). Combined molecular
analysis of LSU and ITS genes places T. chiangmaiensis and
T. cylindrothecia (BCC 3559) within Tubeufia sensu stricto
(Clade F) with strong support (100 % BS and 1.00 PP, Fig. 2).
Other accepted genera
Acanthohelicospora Boonmee & K.D. Hyde gen. nov.,
Index Fungorum number: IF 550572, Facesoffunginumber:
FoF 00206
Etymology: From ‘acantho’ referring to the sexual morph
having setae and ‘helicospora’ referring to the helicosporous
conidia of the asexual morph.
Saprobic on dead wood. Sexual state: Ascomata superfi-
cial, solitary, scattered, globose to subglobose, black,
ostiolate, surrounded with black setae that taper to an acute
apex. Peridium composed of several layers, with outer layer of
cells compressed and black, with inner layer comprising
brown cells of textura angularis. Hamathecium comprising
numerous, filiform, septate, branched, hyaline
pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical, ped-
icellate, with thick, rounded apex, lacking an ocular chamber.
Ascospores overlapping fasciculate, long fusiform-cylindrical,
straight to slightly curved, greater than 7-septate, not constrict-
ed at any septum, hyaline, smooth-walled. Asexual state:
hyphomycetous, helicosporous, helicomyces-like.
Conidiophores mononematous, erect, pale brown to brown,
smooth-walled. Conidiogenous cells holoblastic, terminal or
intercalary, dentate, smooth, with a thickened and truncate
conidiogenous loci. Conidia helicosporous, with wide fila-
ments, coiled, curved, elongated, tapering to narrowly round-
ed ends, multiseptate, not constricted at septum, hyaline,
guttulate, smooth-walled.
Type species: Acanthohelicospora pinicola Boonmee &
K.D. Hyde
Notes: The genus Acanthohe licosporium
has well-
developed spectacular setae on the ascomata, cylindrical asci
and oblong ascospores. The asexual state of
Acanthohelicospora pinicola developed in culture and ap-
pears similar to Helicosporium, except that the conidiophores
are micronematous and not setiferous as in the former (Ellis
1971; Seifert et al. 20 11). Thus, the monotypic genus
Acanthohelicospora is typified by A. pinicola which differs
from all hitherto described species of Acanthostigma and
Acanthostigmina (Réblová and Barr 2000; Kodsueb et al.
2004; Promputtha and Miller 2010; Sánchez and Bianchinotti
2010; Sánchez et al. 2012). Acanthostigma scopulum Peck
and Acanthohelicospora pinicola share some morphological
features (Réblová and Barr 20 00; Kodsueb et al. 2004;
Promputtha and Miller 2010), however, they differ in the
dimensions of the ascomata, asci and ascospores including
the number of septa. Acanthohelicospor a pinicola groups
with an asexual species na med Helicosporium guianense
Linder with high support (96 % BS and 0.99 PP), in a sister
group to Acanthostigma scopulum and Helicosporium aureum
(Corda) Linder with 83 % BS support (Clade K, Fig. 2).
Putatively named strains of Ac anthostigma scopulum, A.
aureum and A. guianense in GenBank are here renamed as
Acanthohelicospora in the phylogenetic tree (Fig. 2), but are
not formally synomymised as these are not ex-type strains and
their characters are unknown.
Acanthohelicospora pinicola Boonmee & K.D. Hyde, sp.
nov., Index Fungorum number: IF 550573,
Facesoffunginumber: FoF 00173 Figs. 5–6
Etymology: Epithet named for the host, Pinus.
Holotype:MFLU10–0049
Saprobic on dead wood of Pinus. Sexual state: Ascomata
116–148μ
m high×108–138μm diam. (x¼ 129:5 122μm ),
superficial, solitary, scattered, globose to subglobose, black,
ostiolate, surrounded by 39–72×5–7.5μm, black setae which
taper to an acute apex. Peridium 12–18.5μm wide, composed
of several layers, outer layer of compressed and black-walled
cells, inner layer comprising brown-walled cells of textura
angularis. Hamathecium comprising numerous, ca.2μm
wide, filiform, septate, branched, hyaline pseudoparaphyses.
Asci 131.5–183×14–18μm (
x¼ 150 16μm ), 8-spored,
bitunicate, cylindrical, pedicellate, with thick and rounded
apex, lacking an ocular chamber. Ascospores 41.5–57×3–
4μm (
x¼ 50:5 3μm ), overlapping fasciculate, long fusi-
form-cylindrical, straight to slightly curved, 7–8(−9)-septate,
not constricted at any septum, hyaline, smooth-walled. Asex-
ual s tate: hyphomyceto us, helicosporous . Conidiophores
mononematous, erect, 3–5μm long×2–4μm wide, pale brown
to brown, smooth-walled. Conidiogenous cells holoblastic,
terminal or intercal ary, dentate, smooth, with a thick-
ened and truncate conidiogenous loci. Conidia
helicosporous, (53-)65–92(−99) μm long×16–20(− 25)
μm diam., filaments 2μm wide, whe n coi led, c urved
andcoiledin1–2 dimension, elongate, tapering to nar-
rowly rounded ends, multiseptate, not constricted at the
septum, hyaline, g uttulate, smooth-walled.
Cultural characteristics: Ascospores germinating on MEA
within 12 h and germ tubes produced at both ends. Colonies
slow growing on MEA, reaching less than 5 mm in 7 days at
28 C, slightly convex, undulating to raised, dentate, with
slightly radial striations and lobate edges, brown. Asexual
state produced in culture. Mycelium superficial, composed of
branched, septate, smooth, hyaline, pale brown to brown
hyphae.
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Fig. 5 Acanthohelicospora pinicola (MFLU 10–0049, holotype). a
Superficial ascomata on substrate. Note ascomata surrounded by black
setae. b L.S. of ascoma. c Peridium with dark setae. d Single seta. e
Pseudoparaphyses. f, g Asci. h–m Ascospores. Scale bars: a–b =100μm,
c–d, h–m =20μm, e =5μm, f–g =50μm
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Material examined: THAILAND, Chiang Rai, Muang, Doi
Tung, elev. ca.1,509mls.,ondeadwoodofPinus L.
(Pinaceae), 6 November 2009, Saranyaphat Boonmee DT-
06 (MFLU 10–0049, holotype = PDD 104451, isotype), ex-
type cultures = MFLUCC 10–0116 = BCC 52036 = IFRD
2196.
Acanthophiobolus Berl., Atti Congl. Bot. Intern. Di Ge-
nova, 1892: 571 (1893) [1892]
Saprobic on cloth. Sexual state: Ascomata superficial,
globose to subglobose, reddish-brown to dark brown, ostiole
central, with red brown to dark brown setae, setae septate,
tapering towards apex. Peridium comprising several-layers;
inner layer composed of textura prismatica-porrecta and hy-
aline to pale, outer layer composed of 2–3 layers of dark
brown cells of textura angularis. Hamathecium comprising
filiform pseudoparaphyses. Asci 8-spored, bitunicate, elon-
gate, cylindro-clavate, short pedicellate. Ascospores spirally
arranged in ascus, filiform, septate, hyaline (from Boonmee
et al. 2011). Asexual state:Unknown.
Notes: This monotypic genus for which no molecular data
are available was illustrated in Boonmee et al. (2011). The
genus differs from Acanthostigma morphologically. The
ascomata of Acanthophiobolus are bright coloured with
much longer setae, and the asci are elongate-cylindrical
with long filiform, trans-septate ascospores. We there-
fore maintain Acanthophiobolus as a distinct genus in
Tubeufiaceae.
Type species: Acanthophiobolus helminthosporus (Rehm)
Berl., Die Pilze des Weinstockes, Vienna: 571 (1893) [1892]
Acanthostigma De Not., Sfer. Ital., 85 (1863)
Saprobic on dead wood. Sexual state: Ascomata superfi-
cial, scattered, globose to subglobose, mostly dark brown to
black, occasionally collapsing when dry, with distinct dark
setae, ostiolate. Peridium consisting of several-layers of
Fig. 6 Acanthohelicospora pinicola (MFLU 10–0049, holotype). Col-
onies and asexual state on MEA culture. a Germinating ascospore. b, c
Colonies on MEA from surface and reverse. Note dark brown colonies. d
Arial mycelium on culture. e–g Conidiophores developing on hyphae. h
Conidiogenous cells. i–l Conidia. Scale bars: a–c = 10 mm, d–i =5μm,
e–l =10μm
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textura angularis. Hamathecium with numerous, cellular,
branching and anastomosing pseudoparaphyses. Asci 8-
spored, bitunicate, fissitunicate, clavate, short pedicellate,
broadly rounded and thickened at apex. Ascospores overlap-
ping 2–3-seriate, fusiform, narrowly rounded at both ends,
with one of middle cells often broader than others, tapering
towards the ends, trans-septate, straight or slightly curved,
not-constricted or slightly constricted at septa, mostly hyaline,
guttulate when immature, smooth-walled. Asexual state :
Chlamydospores only known from culture (from Boonmee
et al. 2011).
Notes: Acanthostigma sensu stricto is based on
Acanthostigma perpusillum and the description given by de
Notaris (1863). Réblová and Barr (2000) monographed and
accepted six species in Acanthostigma. Morphologically all
species are characterised by dark brown to black ascomata
covered by dark setae; ascospores are usually broadly fusi-
form to clavate, asymmetrical, trans-septate and hyaline. The
multigene phylogenetic analysis indi cates that several
Acanthostigma epithets are polyphyletic and belong within
the Tubeufiaceae (Promputtha and Miller 2010; Boonmee
et al. 2011; Sánchez et al. 2012).
The type species A. perpusillum was described and illus-
trated in Boonmee et al. (2011), while a new species,
A. chiangmaiensis Boonmee & K.D. Hyde, was also intro-
duced. Other species previously regarded as Acanthostigma
(e.g. Acanthostigma filiforme Promp. & A.N. Mill.,
A. minutum (Fuckel) Sacc.) are scattered in the phylogenetic
tree, indicating that ascomata with setae have evolved on more
than one occasion and is not a reliable character for determin-
ing genera in this family. We searched for herbarium speci-
mens of Acanthostigma perpusillum from several herbaria
(e.g. GE, PAD, RO and TO), but none are available for study.
Réblová and Barr (2000) reexam ined, described and
illustrated this species and Boonmee et al. (2011)madedraw-
ings of the taxon based on Réblová and Barr (2000). The
species has brown opaque, short setae on reddish-brown to
dark brown ascomata. In this paper we treat Acanthostigma
perpusillum strain UAMH 7237 as an authentic specimen to
represent Acanthostigma sensu stricto. Molecular analysis
indicat that A. perpusillum and A. chiangmaiensis has moder-
ate support (72 % BS) in Clade E (Fig. 2). A putative strain of
A. minutum (MVB 781 (BBB), Argentina, Sánchez et al.
2012) forms a long subbranch with A. perpusillum
and
A. chiangmaiensis but with weak support. Therefore, two
species A. chiangmaiensis and A. minutum are accepted in
the genus based on phylogeny (Fig. 2). Helicosporium sensu
stricto forms a sister clade (Clade D). No asexual state has
been reported for Acanthostigm a perpusillum, however,
A. chiangmaiensis formed chlamydospores in culture. Addi-
tionally A. filiforme Promp. & A.N. Mill. and
A. septoconstrictum Promp. & A.N. Mill. and
A. multiseptatum described in Promputtha and Miller (2010)
are transferred to Neoacanthostigma and Acanthostigmiana
respectively based on morphology and molecular data.
Type species: Acanthostigma perpusillum de Not., Sfer.
Ital.: 207 (1863)
Acanthostigmina Höhn., Sber. Akad. Wiss. Wien, Math.-
naturw. Kl., Abt. 1 118: 1499 [39 repr.] (1909),
Facesoffunginumber: FoF 00207
Saprobic on rotten wood, widespread in tropical regions.
Sexual state: Ascomata superficial, solitary, scattered, some
clustered, globose to subg lobose, dark brown, ostiolate,
surrounded by sparse, dark brown to black setae; setae rela-
tively long, tapering to an acute apex. Peridium thick, com-
prising several layers of textura angularis to subglobosa,
inwardly comprising brown to reddish-brown, small brown
cells of textura subprismatica. Hamathecium comprising nu-
merous, filiform, septate, branched, hyaline and pale brown
pseudoparaphyses, embedded in a gelatinous matrix. Asci 8-
spored, bitunicate, broadly cylindric-subclavate, pedicellate,
thickened at apex, without an ocular chamber. Ascospor es 2–
3-seriate, elongate, cylindric-fusiform, curved, tapering to-
wards sub-rounded ends, less than 10-septate, slightly con-
stricted at septum, hyaline when young and pale brown at
maturity, some surrounded by a mucilaginous sheath, smooth-
walled. Asexual state: Unknown.
Type species: Acanthostigmina minuta (Fuckel) Clem. &
Shear
Notes: The genus Acanthostigmina was introduced by von
Höhnel (1909a)withA. minuta (Fuckel) Höhn. as the type
species. Saccardo (1883) had earlier included this species in
Acanthostigma. Subsequently Acanthostigmina minuta has
been placed in other genera (Barr 1980; Crane et al.
1998;
Réblová and Barr 2000). In this study, we re-examined a
specimen of Acanthostigmina minutum from Germany which
Réblová and Barr (2000)consideredtoberepresentativeof
the species (Rehm exsiccatae, Ascomycete No. 1568, PAD)
(Fig. 7). Rehm’s exsiccatae specimen (BPI 624355) is char-
acterized by dark brown to black ascomata, moderately stiff
setae, broadly cylindric-subclavate, apically thickened asci
and 5–9-septate, hyaline to pale brown ascospores with a thin
mucilaginous sheath. Thi s differs from the description of
Acanthostigma minutum from the PAD neotype provided by
Réblová and Barr (2000) in which the ascospores were
more than 10-septate, hyaline and without a mucilagi-
nous sheath, whereas in other characters t hey are rather
similar. Unlike Ac ant hos tigm in a minut a, Acanthostigma
perpusillum has ascomata covered by numerous setae; in
addition these species differ in the size and shape of
asci and ascospores.
Molecular data indicate that three non-type isolates of
Acanthostigmina minutum from North America
(ANM283, ANM818 and ANM880) formed a single
clade distant from the Acanthostigma perpusillum clade
(Promputtha and Miller 20 1 0). Sánchez and Bianchinotti
254
Fungal Diversity (2014) 68:239–298
( 2010) found a collection in Argentina identified as
Acanthostigma minutum MVB 781 (BBB), which
Sánchez et al. (2012)showedtoclusterwith
Acanthostigma perpusillum. In our multigene phyloge-
netic analysis, the North American strains of
Acanthostigmina minuta (ANM810, ANM818 and
ANM238) formed a monophyletic clade that clustered
with two strains of Acanthostigmina multiseptatum
(ANM475 and ANM665), although with weak support
(Clade L, Fig. 2). We therefore treat Clade L as
Acanthostigmina sensu stricto comprising A. minuta
and A. multiseptat um.
Fig. 7 Acanthostigmina minuta (BPI 624355). a Ascomata. b LS of ascoma. c Setae. d Peridium. e Pseudoparaphyses. f, g Asci. h, i Ascospores. Scale
bars: a–b =200μm, c–d, f–i =50μm, e =5μm
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Fungal Diversity (2014) 68:239–298
Type species: Acanthostigmina minuta (Fuckel) Clem. &
Shear, Gen. Fung. (Minneapolis): 270 (1931),
Facesoffunginumber: FoF 00174 Fig. 7
≡ Lasiosphaeria minuta Fuckel, Jb. nassau. Ver. Naturk.
23–24: 148 (1870) [1869–70]
Saprobic on decaying wood of a rotten branch of Fagus
sylvatica. Sexual state: Ascomata (211-)234–299μm
high×(216-)226–270μm diam., superficial, solitary, scattered,
occasionally clustered, globose-subglobos e, dark brown,
ostiolate, surrounded by sparse, erect, dark brown to black
setae, (30-)56–84(−105) μm long, tapering towards an acute
end at apex. Peridium 29–34μm wide, with several layers of
brown to reddish-brown cells, comprising textura angularis to
subglobosa, inwardly with small brown cells of textura
subprismatica. Hamathecium comprising ca. (1-)1.5–2μm
wide, numerous filiform, septate, branched, hyaline to pale
brown pseudoparaphyses, embedded in a gelatinous matrix.
Asci (73-)84–118×20–29.5μm (
x¼ 98 24μm , n=20), 8-
spored, bitunicate, broadly cylindric-subclavate, pedicellate,
thickened at apex, without an ocular chamber. Ascospores
(42-)47–60×6–9μm (
x¼ 52 7μm , n =20), 2–3-seriate,
elongate-fusiform, curved, tapering towards rounded ends,
(6-)7–9-septate, broader at fourth and fifth cells from apex,
slightly constricted at septum, hyaline to pale brown when
mature, occasionally surrounded by a mucilaginous sheath,
smooth-walled. Asexual state: Unknown.
Material examined: GERMANY, Bavarian Alps M ts.,
Kampenwand, elev. ca 1200 msl., on decaying wood branch
of Fagus sylvatica L. (Fagaceae), June 1904, Rehm H.,
Ascomyceten no. 1568 (BPI 624355).
New combination:
Acanthostigmina multiseptatum (Promp. & A.N. Mill.)
Boonmee & K.D Hyde com. nov., Index Fungorum number:
IF 550681
≡ Acanthostigma multiseptatum Promp. & A.N. Mill.,
Mycologia 102(3): 577 (2010)
Aquaphila Goh, K.D. Hyde & W.H. Ho, Mycol. Res.
102(5): 588 (1998)
Saprobic on submerged woody substrates, widespread in
tropical regions. Mycelial colonies partly developed inside
wood and partly on wood surface. Sexual state: Ascomata
superficial, gregarious, globose to subglobose, dark brown to
black, with brown setae, straight, thick-walled, tapering to an
acute apex. Peridium outer layer of textura angularis com-
prising black-walled cells, cell of inner layer, elongate, pale
brown to subhyaline. Pseudoparaphyses filiform, ca. 2mm
wide, hyphae-like, branc hed, hyaline. Asci 8-spored,
bitunicate, cylindroclavate, with long pedicels, apically round-
ed. Ascospores 2–3-seriate, cylindrical fusiform, 7-transverse
septa, hyaline to pale brown, smooth (from Tsui et al. 2007).
Asexual state: Conidiophores semi-macronematous,
mononematous, borne as lateral branches from superficial
hyphae, hy aline, delicate, septate, simple or br anched,
flexuous to geniculate. Conidiogenous cells integrated, termi-
nal or intercalary, denticulate, monoblastic or polyblastic,
proliferation sympodial, indeterminate. Conidial secession
schizolytic. Conidia borne singly, acrogenous and becoming
lateral as a result of conidiophore proliferation, hyaline, fusoid
to falcate or sigmoid, br oad, multi-euseptate, apedicellate
(from Goh et al. 1998).
Notes
: Clade J represents Aquaphila and comprises species
with darkly pigmented ascomata covered by dark setae,
cylindrical-clavate and apically thickened asci, and fusiform
6–7-septate ascospores. The asexual form occurs in an aquatic
habitat and produces curved conidia that are distinct from the
type species in the sister genus Chlamydotubeufia,
C. huaikangplaensis that produces dictyochlamydospores
(Tsui et al. 2007; Boonmee et al. 2011) and other Tubeufia
species. The ex-type strain of Tubeufia asiana (BCC 3463)
clusters in Clade J with Aquaphila albicans Goh, K.D. Hyde
& W.H. Ho, with robust support (100 % BS and 1.00 PP).
Tubeufia asiana was introduced as the asexual state of
Aquaphila albicans by Tsui et al. (2007). Our phylogenetic
analysis shows it to clusters far from Tubeufia sensu stricto;
Aquaphila albicans is the older name and thus Tubeufia
asiana is a synonym.
Type species: Aquaphila albicans Goh, K.D. Hyde & W.H.
Ho, Mycol. Res. 102(5): 588 (1998)
≡ Tubeufia asiana Sivichai & K.M. Tsui, Mycologia 99(6):
885 (2008) [2007]
Bifrontia Norman, Bot. Notiser: 18 (1872),
Facesoffunginumber: FoF 00208
Saprobic on dead wood. Sexual state: Ascomata superfi-
cial, seated on a subiculum, solitary or gregarious, globose to
subglobose, black, cupulate, turbinate, collapsing when ma-
ture or dried, central ostiole. Peridium thick-walled, outer
layer composed of 3–4-layers of brown cells of textura
subangularis, innermost layers of 2–3-layers of hyaline cells
of textura subprismatica. Hamathecium with sparse
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, fu-
siform, broadly rounded above, thick-walled, narrowing to-
wards the base, short-pedicellate, arranged in rows on basal
cells of ascomata. Ascospores overlapping fasciculate, long-
fusiform, straight to slightly curved, 7–9-septate, constricted
at septa, pale brown, smooth-walled. Asexual state:
Unknown.
Lectotype species designated here: Bifrontia compactior
Norman, Bot. Notiser: 19 (1872), Facesoffunginumber:FoF
00175 Fig. 8
Fig. 8 Bifrontia compactior (O-F72543, holotype). a, b Herbarium
material and labels. c, d Black ascomata on host surface. e Vertical
section of ascoma. f, g Close up of peridium and sparse
pseudoparaphyses. h-j, l, m Asci with ascospores - note l, m,stainedin
cotton blue reagent; k, n-p Ascospores; note n stained in cotton blue
reagent. Scale bars: c =200μm, d =100μm, e =50μm, f =20μm, g =
10μm, h–j, l–m =10μm, k, n–p =5μm
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Saprobic on dead wood of Salix glauca. Sexual state:
Ascomata 100–150μm high×190–250μm diam., superficial,
seated on a subiculum, solitary or gregarious, globose to
subglobose, black, cupulate, turbinate, collapsing when ma-
ture or dried, central ostiole ambiguous. Peridium thick-
walled, 35–50μm wide, composed of 3–4-layers of brown
cells of textura angularis, innermost layers of 2–3-layers of
hyaline cells of textura subprismatica. Hamathecium com-
prising sparse pseudoparaphyses. Asci 55–100×14.5–25μm
(
x¼ 75 17μm , n=20), 8-spored, bitunicate, fissitunicate,
fusiform, broadly rounded above, thick-walled, narrow to-
wards at base, short-pedicellate, arranged in rows on basal
cells of ascomata. Ascospores 40–60×4–6μm (
x¼ 48 5μm ,
n=10), overlapping fasciculate, long-fusif orm, straight to
slightly curved, 7–9-septate, constricted at septa, pale brown,
smooth-walled. Asexual state: Unknown.
Material examined: NORWAY, Finnmark, Hammerfest:
Prope Hammerfest Finmarki , on wood of Salix glauca L.
(Salicaceae), J.M. Norman, date of collection unknown (O-
F72543, holotype).
Notes: Bifrontia was introduced by Norman (1872)with
two species but neither was designated as the type for the
genus. Bifrontia compactior Norman was listed first in the
diagnosis for the genus and i s typical of the protologue.
Therefore we designate herein B. compactior as the lectotype
of the genus. This species has t ypical characte ristics of
Tubufiaceae except that the hamathecium rarely includes
pseudoparaphyses. We refer this genus to T ubeufiaceae.
Boerlagiomyce s Butzin., Willdenowia 8(1): 39 (1977),
Facesoffunginumber: FoF 00209
Boerlagella Penz. & Sacc., Malpighia 11(9–10): 404
(1897)
Saprobic on decaying wood. Sexual state: Ascomata su-
perficial, seated on a subiculum, globose, scattered, with
flexuous, hairy, black, velvety, septate setae, surrounded by
black mycelium. Peridium composed of carbonaceous, multi-
layered walls. Hamathecium composed of filiform, hyaline
pseudoparaphyses. Asci 8-spored, bitunicate, cylindric-cla-
vate, narrowing towards base, short-pedicellate or
apedicellate, apically rounded. Ascospores 2–3-seriate, cylin-
dric-fusoid, gently curved, obtuse at both ends, muriform,
multiseptate, not constricted at septa, hyaline, smooth-walled
(adapted from Penzig and Saccardo 1904). Asexual state:
Unknown.
Notes: The genus Boerlagiomyces was introduced by
Butzin (1977)withB. velutinus (Penz. & Sacc.) Butzin as
the type species. The genus is distinctive in producing super-
ficial, whitish to pale brown, soft ascomata with large hyaline,
muriform ascospores that often develop on submerged wood
in freshwater. Crane et al. (1998) monographed and accepted
six species in Boerlagiomyces in
Tubeufiaceae.Thisgenus
presently com prises nine species acc ording to Index
Fungorum (2014 ). A putative strain of Boerlagiomyces
websteri Shearer & J.L. Crane was placed with Rhytisma
acerinum (Pers.) Fr. in Rhytismataceae by Kodsueb et al.
(2006); this seems unlikely. Garethjonesia has been consid-
ered a synonym of Boerlagiomyces (Stanley and Hyde 1997;
Crane et al. 1998), however they are morphologically distinct.
The habitats (aquatic versus terrestrial) also differ. We treat
Garethjonesia here as a good genus in Ascomycetes incertae
sedis.
Type species: Boerlagiomyces velutinus (Penz. & Sacc.)
Butzin, Willdenowia 8(1): 39 (1977), Facesoffunginumber:
FoF 00176 Fig. 9
≡ Boerlagella velutina Penz. & Sacc., Malpighia 11(9–10):
404 (1897)
Saprobic on decaying wood or culms in petioles of
Plectocomia sp. Sexual state: Ascomata 500–600μm diam.,
superficial, seated on black subiculum, globose to pyriform,
dark brown, scattered, ostiolate, with sparse, flexuous, black,
septate, velvety, hyphal appendages, 210–25 0×3–6μm,
black, septate, surrounded black mycelium. Peridium com-
posed of carbonaceous, multilayered walls. Hamathecium
composed of filiform, hyaline pseudoparaphyses. Asci 200–
250×30–35μm, 8-spored, bitunicate, elongate, cylindric-cla-
vate, apically rounded, narrowing towards base, short-
pedicellate or apedicellate. Ascospores 90–120×12–14μm,
2–3-seriate, cylindric-fusoid, gently curved, obtuse at both
ends, muriform, 25–
30-septate, not constricted at septa, hya-
line, smooth-walled (adapted from Penzig and Saccardo
1904). Asexual state: Unknown.
Chlamydotubeufia Boonmee & K.D. Hyde, Fungal Diver-
sity 51(1): 78 (2011)
Saprobic on decaying wood in terrestrial habitats, wide-
spread in temperate to trop ical regio ns. Sexual state:
Ascomata superficial, solitary, globose-subglobose, ostiolate,
surrounded by dark setae. Peridium thick-walled, dark brown
to black, composed of cells of textura angularis.
Hamathecium composed of cellular hyaline
pseudoparaphyses, embedded in a mucilaginous matrix. Asci
8-spored, bitunicate, fissitunicate, cylindric-clavate to broadly
clavate, short-pedicellate, rounded at apex. Ascospores 2–3-
seriate, hyaline, narrowly fusiform, broad at supra-median,
slightly curved, multiseptate, slightly constricted at septum,
with asym metrical ends. Asexual state: h yphomycetous,
helicosporous, also producing a dictyochlamydosporous state
in culture and often on wood. Dictyochlamydospores broadly
oblong, elongate, multiseptate, at first reddish-brown, becom-
ing black (from Boonmee et al. 2011).
Type species: Chlamydotubeufia huaikangplaensis
Boonmee & K.D. Hyde, Fungal Diversity 51(1): 78 (2011)
Figs. 10–11
Chlamydotubeufia (Clade I) includes terrestrial taxa and
produces dictyochlamydospores. This genus is distinct from
the aquatic genus Aquaphila (Clade J) which produces sickle-
shaped conidia (Tsui et al. 2007; Boonmee et al. 2011). In
258
Fungal Diversity (2014) 68:239–298
addition to the type species, C. huaikangplaensis, two other
species are included in this genus, specifically
C. chlamydosporum and C. khunkornensis. A new collection
of Chlamydotubeufia huaikangplaensis is illustrated below
(THAILAND, Chiang Mai, Mae Taeng, Mushroom Research
Centre, N19°17.123′ E98°44.009′,elev.ca. 900 msl., on
dead wood of an unidentified tree, 23 June 2011, Saranyaphat
Boonmee, MRC-02 (MFLU 11–1145) = living culture =
MFLUCC 11–0509).
Helicangiospora Boonmee, Bhat & K.D. Hyde, gen. nov.,
Index Fungorum number: IF 550574, Facesoffunginumber:
FoF 00210
Etymology: Helicangiospora, referring to conidial spores
produced endogenously within a vessel-like conidiogenous
cell.
Saprobic on dead wood. Sexual state: Ascomata superfi-
cial, solitary to clustered, scattered, globose to subglobose,
dark brown to black, with abundant setae, ostiolate. Setae stiff,
tapering to an acute tip at apex. Peridium composed of several
outer layer cells of textura angularis, darkened, innermost
layer comprising pale brown to hyaline cells of textura
subprismatica. Hamathecium comprising ca. 2μm wide, nu-
merous, filiform, septate, branched, hyaline
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,
cylindrical-clavate or saccate, with rounded apex, pedicellate,
ocular chamber not apparent. Ascospores overlapping 1–3-
seriate, fusiform, hyaline, tapering toward the ends,
supramedianly wider, trans-septate, smooth-walled. Asexual
state: hyphomycetous, helicosporous. Conidiogenous cells
terminal, integrated, holoblastic, globose, occasionally
Fig. 9 Boerlagiomyces velutinus
(Redrawn from Penzig and
Saccardo 1904), as Boerlagella
velutina. a, b Substrate and
ascoma, 500–600μm diam. c
Ascus, mycelium and
pseudoparaphyses, ascus size:
200–250×30–35μm d
Ascospores, 90–120×12–14μm
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subglobose to oval, golden-brown to olive-brown. Conidia
helicosporous, developing endogenously within
conidiogenous cells, released by breaking wall of
conidiogenous cell, trans-septate, tapering towards rounded
ends, not constricted at septum, coiled once, hyaline, pale
brown at maturity.
Type species: Helicangiospora lignicola Boonmee, Bhat &
K.D. Hyde
Fig. 10 Chlamydotubeufia huaikangplaensis (MFLU 11–1145). a Ascomata on substrate. b L.S. of ascoma. c Single seta. d L.S. of peridium. e
Pseudoparaphyses. f–h Asci. i–k Ascospores. Scale bars: a, b =100μm, c, f–h =50μm, d =40μm, e =5μm, i–k =20μm
260
Fungal Diversity (2014) 68:239–298
Notes: Phylogenetically Helicangiospora lignicola does
not group with any other taxa and is basal to Clades A-B,
although lacks support. The genus is also morphologically
distinct as it produces a unique helicoma-like asexual state
in culture (Fig. 13). The holoblastic conidia are initiated at the
tip of a conidiogenous cell by formation of a globose, moder-
ately dark brown cell within which t he characteristic
helicosporous conidia differentiate endogenously. The heli-
coid conidia are released by schizolytic break down of the
outer thick-walled sphaerical capsule. Endogenous differenti-
ation of the conidium is seen in most phialidic type
conidiogenesis wherein the conidial primordium is differenti-
ated enteroblastically within the phialide, but in most genera
maturation of conidia is exogenous. In asexual genera, such
as, Chalara, Fusichalara, Sporoschisma, and
Sporoschismopsis, not only differentiation but also maturation
of the conidia takes place within the venter of the phialide
(Bhat 2010). However, these structures are not similar to what
is seen in the asexual state of the present specimen. Therefore,
the genus Helicangiospora is established to accommodate this
unique species that produces helicosporous conidia within a
conidiogenous capsule.
Fig. 11 Chlamydotubeufia huaikangplaensis (MFLU 1 1–1145). Colo-
nies on MEA. a Germinating ascospore. b, c Colonies on MEA from
surface and reverse. Note dark brown colonies. d Chlamydospores
growing on substrate. Note chlamydospores produced on media, with
and without substrate. e Chlamydospores on plant tissues. f–j Chlamydo-
spores. Scale bars: a =20μm, b–d = 10 mm, e–j =40μm
261
Fungal Diversity (2014) 68:239–298
Type species: Helicangiospora lignicola Boonmee, Bhat &
K.D. Hyde, sp. nov., Index Fungorum number: IF 550575,
Facesoffunginumber: FoF 00177 Figs. 12–13
Etymology:Fromlignin referring to wood and -icola mean-
ing associated with in reference to growing on wood.
Holotype:MFLU11–0137
Saprobic on dead wood . Sexual state: Ascomata
(216-)296– 351μ m high×266– 340μ m diam.
(
x¼ 286 308μm ), superficial, solitary to clustered,
scattered, globose to subglobose, dark brown to black, with
abundant setae, ostiolate. Setae stiff, tapering to an acute tip at
apex, 56–77μm long. Peridium 36–56μm wide, composed of
several outer cells of textura angularis, darkened; innermost
comprising thin layers of pale brown to hyaline cells of textura
subprismatica. Hamathecium comprising ca. 2μm wide, nu-
merous, filiform, septate, branched, hyaline
pseudoparaphyses. Asci (77-)80– 106 ×15– 19 μ m
(
x¼ 93 17μm , n=20), 8-spored, bitunicate, fissitunicate,
Fig. 12 Helicangiospora lignicola (MFLU 11–0137, holotype). a Ascomata on substrate. b L.S. of ascoma. c L.S. of peridium. d Single seta. e
Pseudoparaphyses. f–i Asci. j–l Ascospores. Scale bars: a–b =100μm, c, j–l =20μm, d =40μm, e =5μm, f–i =50μm
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Fungal Diversity (2014) 68:239–298
cylindrical-clavate or saccate, with rounded apex, pedicel ca.
11μm long, ocular chamber not apparent. Ascospores
(40-)45–53×5–8μm (x¼ 47 7μm , n=20), overlapping 1–
3-seriate, fusiform, hyaline, slightly curved, tapering toward
Fig. 13 Helicangiospora lignicola (MFLU 11–0137, holotype). a Ger-
minating ascospore. b, c Colonies on MEA from surface and reverse. d, e
Mycelium and development of conidia in culture. f Conidiogenous cells.
Note the formation of conidium at this stage. g–j Endogenous develop-
ment of conidium. j Release of conidium. k–n Conidia. Scale bars: a =
20μm, b, c =10mm,d–n =5μm
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ends, supramedianly wider, 6–7-septate, not constricted at
septum, smooth-walled. Asexual state: hyphomycetous,
helicosporous. Conidiogenous cells terminal, integrated, ho-
loblastic, globose, sometimes subglobose to oval, golden-
brown to olive-brown, 12– 17.5 μ m diam. Conidia
helicosporous, differentiating endogenously within
idiogenous cells, rele ased by break down of wall of
conidiogenous cell, 12–17μm diam when coiled, filaments
4–7μm wide, 5-septate, tapering towards rounded ends, not
constricted at septum, coiled once, hyaline, pale brown at
maturity, mostly smooth-walled, sometimes verrucose, slight-
ly bulging at the septa.
Cultural characteristics: Ascospores germinating on MEA
within 12 h. Colonies slow growing on MEA, less than 5 mm
diam in 7 days at 28 C, slightly convex, with an undulate edge,
white or pale brown and dark brown, white at margin.
Mycelium superficial, septate, branched, smooth, pale brown,
with hyphae producing erect, septate conidiophores.
Material examined: THAILAND, Chiang Rai, Muang, Doi
Pui, elev. ca.403–936 msl., on dead wood of an unidentified
tree, 10 May 2011, Saranyaphat Boonmee, DP-03 (MFLU
11–0137 holotype; PDD 104452 isotype); ex-type living
culture = MFLUCC 11–0378 = BCC 52029 = ICMP 20069.
Helicoma Corda, Icon. fun g. (Pragu e) 1: 1 5 (1837),
Facesoffunginumber: FoF 00211
Possible synonyms (Index Fungorum 2014, not seen)
Lituaria Riess, Bot. Ztg. 11: 136 (1853)
Helicocoryne Corda 1854, Icon. fung. (Prague) 6: 9 (1854)
Drepanospora Berk. & M.A. Curtis [as ‘Drepanispora’], in
Berkeley, Grevillea 3(no. 27): 105 (1875)
Helicopsis P. Karst., Revue mycol., Toulouse 11(no. 42):
96 (1889)
Troposporella P. Karst., Hedwigia 31: 299 (1892)
Helicosporella G. Arnaud, Bull. trimest. Soc. mycol. Fr.
69: 292 (1954) [1953]
Helicosporina G. Arnaud ex Rambelli, Mycopath. Mycol.
appl. 13: 110 (1960)
Helicominopsis Deighton, Mycol. Pap. 78: 20 (1960)
Moorella P.Rag. Rao & D. Rao, Mycopath. Mycol. appl.
22: 51 (1964)
Saprobic on woody substrata. Mycelium partly immersed,
pale brown, septate, branched hyphae. Sexual state: Ascomata
superficial, solitary, sometimes clustered, gregarious, superfi-
cial, subglobose, oval to obovoid, with soft texture, dark
brown, pale brown above, with a central papillate ostiole,
rarely with setae.
Setae up to 80μm long, dark brown, tapering
towards subacute apex, widest at base, septate. Peridium thick-
walled, composed of brown to reddish-brown cells of textura
angularis. Hamathecium comprising numerous, filiform, sep-
tate, branched, hyaline pseudoparaphyses. Asc i 8-spored,
bitunicate, elongate to cylindric-clavate, thickened at apex,
pedicellate. Ascospores 2–3-seriate, elongate-fusiform,
subcylindrical, tapering towards subacute ends, up to 10-
septate, not constricted at septum, hyaline, smooth-walled.
Asexual state: Conidiophores superficial, macronematous,
crowded, erect, dark brown, septate, rarely branched, darkened
and slightly constricted at septum. Conidiogenous cells
monoblastic to polyb lastic, sometimes branched at apex,
browntodarkbrown,smooth.Conidia tightly coiled 1–1½
times, conidial filament 6–9μm wide, hyaline to pale brown,
tapering towards the flattened end with a basal scar, septate,
slightly constricted at septum, smooth-walled.
Type species: Helicoma muelleri Corda
Notes: The genus Helicoma was introduced by Corda
(1837), with the type species H. muelleri Corda. The genus
is distinguished by its relatively short, erect, thick, dark
brown, smooth conidiophores, holoblastic coni diogeno us
cells and helicoid, hyaline, thick-walled, brown to dark brown
conidia forming from terminal, denticulate conidiophores
(Goos 1986; Seifert et al. 2011). Illustrations presented here
are based on BPI 447569 considered authentic material of
H. muelleri on natural substrate and determined by D. Linder.
Variou s Helicoma species have been connected with sexual
taxa in Tubeufiaceae and the links are confused (Tsui et al.
2006, 2007;Zhaoetal.2007; Boonmee et al. 2011). The
sexual state is described from Helicoma rugosa (BPI
1104599, dried culture: BER 12 80–4). Putatively named
strains of Helicosporium linderi R.T. Moore (NBRC 9207),
T. paludosa (P. Crouan & H. Crouan) Rossman (HKUCC
9118, ANM 196, ANM 1169) and Thaxteriella helicoma
(W. Philli ps & Plowr.) J.L. Crane, Shearer & M.E. Barr
(JCM 2739, UBCF 13877) clustered in this clade and we
rename them as Helicoma in the phylogenetic tree (Fig. 2),
but do not formally synonymize them as we have not seen
voucher material. Asexual morphs regarded as Helicoma
rugosa formed in cultures of Thaxteriella helicoma (JCM
2739 and UBCF 13877). Tubeufia khunkornensis Boonmee
&K.D.Hyde(MFLUCC10–0119, ex-type strain),
T. inthanonensis Boonmee & K.D. Hyde (MFLUCC 11–
0003, ex-type strain) and T. miscanthi W.H. Hsieh, Chi Y.
Chen & Sivan., epitypified here are synonymized under
Helicoma. Helicoma chiangraiense, H. fagacearum and
H. siamense formed distinct groups at the base of Clade A,
and we therefore introduce them as new species of Helicoma.
Type species: Helicoma muelleri Corda, Icon. fung. (Prague)
1: 15 (1837), Facesoffunginumber: FoF 00178 Fig. 14
Saprobic on woody substrates. Mycelium composed of
partly immersed and partly superficial, pale brown, septate,
branched hyphae. Sexual state: Unknown. Asexual state:
Conidiophores (43-)80–153.5μm long×7–10μm wide
(
x¼ 106 8μm , n = 20), superficial, macronematous,
crowded, erect, dark brown, septate, rarely branched, dark-
ened at septum. Conidiogenous cells monoblastic to
polyblastic, sometimes branched at apex, brown to dark
brown, smooth-walled. Conidia (15-)16–19(−21) μm diam.,
and conidial filament 6–9μm wide (
x¼ 17 7μm , n=20),
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tightly coiled 1–1½ times, hyaline to pale brown, tapering
toward flat end and with a basal scar 4–7(−8) μm wide,
septate, slightly constricted at septum, smooth-walled.
Material examined: USA, New Hampshire, Bartlett, on
dead wood (undermined substrate), R. Thaxter, April 1901;
Detr. D. Linder, BPI 447569.
Notes: The sequence (Helicoma muelleri CBS 964.69) of
this species was used in the molecular analysis of Tsui et al.
(2006). We treat this as an authentic specimen but the species
needs epitypfiying with a fresh collection.
New combinations:
Helicoma khunkornense (Boonmee & K.D. Hyde) S.
Boonmee & K.D. Hyde, comb. nov., MycoBank: MB 804554
≡ Tubeufia khunkornensis Boonmee & K.D. Hyde, Fungal
Diversity 51(1): 86 (2011).
Notes: This species produced a helicoma-like asexual state
in culture, with brown helicoid conidia on thick robust conid-
iophores (Fig. 4, in Boonmee et al. 2011). It groupe d in
Helicoma sensu stricto (Clade A) with high support (95 %
BS and 1.00 PP) in the phylogenetic analysis (Fig. 2)and
therefore is synonymised under Helicoma.
Material examined: THAILAND, Chi ang Rai, Muang,
Khun Korn Waterfall, elev. as 671 msl., N19°51 –54′ E
99°35.39′, on dead wood of an unidentified tree, 13 November
2009, S. Boonmee KK-08 (MFLU 10–0052, holotype), ex-
type culture MFLUCC 10–0119 = IFRD 2180 = BCC 52297.
Helicoma inthanonense (Boonmee & K.D. Hyde) S.
Boonmee & K.D. Hyde, comb. nov., MycoBank: MB 804555
≡ Thaxteriella inthanonensis Boonmee & K.D. Hyde, Fun-
gal Diversity 51(1): 86 (2011).
Notes: This species produces a helicoma-like asexual state
in culture, with brown helicoid conidia formed on slender,
hyaline conidiophores (Boonmee et al. 2011). It may also be
similar to Drepanospora pannosa Berk. & M.A. Curtis as it
forms secondary microconidia (Seifert et al. 2011). This was
considered to be the asexual state of Tubeufia helicoma. In the
phylogenetic tree in Clade A (Fig. 2), this appears to be a
different species from Helicoma rugosa (=Tubeufia
helicoma), but all three taxa are congeneric.
Material examined: THAILAND, Chinag Mai, Doi
Inthanon, Jom Thong, elev. 800–1,000 msl., N18°31.576′ E
98°29.790′,ondeadbarkofanunidentifiedtree,16
Fig. 14 Helicoma muelleri (BPI
447569). a Conidiophores with
attached apical conidium on
natural substrate. b, c Squash
mount of conidiophores with
conidial development at the apex
(arrows). d Conidiophores with
detached conidia and minute
denticles (arrows). e–g Conidia.
Scale bars: a, d–g =20μm, b–c =
50μm
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November 2010, Rungtiwa Phookamsak (MFLU 11–0003,
holotype), ex-type culture MFLUCC 11–0003 = BCC 52153.
Helicoma rugosa (C. Booth) S. Boonmee & K.D. Hyde,
comb. nov., Facesoffunginumber: FoF 00179 Fig. 15
≡ Tubeufia rugosa C. Booth, Mycol. Pap. 94: 13 (1964)
≡ Thaxteriella helicoma (W. Phi llips & Plowr.) J.L.
Crane, Shearer & M.E. Barr, Can. J. Bot. 76(4): 610
(1998)
= Sphaeria helicoma W. Phillips & Plowr., Grevillea, 6:26
(1877)
Fig. 15 Helicoma rugosa (BPI 1104599, dried culture: BER 12 80–4). a
Ascomata. b L.S. ascoma. c Single seta. d Peridial w all. e
Pseudoparaphyses. f–h Asci. i–k Ascospores. l Colony on dried culture.
m Mycelium. n Conidiophore (arrows). o–q Conidiophores with at-
tached conidia. Scale bars: a–c =100μm, c–d, i–k, n–q =20μm, f–h =
50μm, e, m =5μm, l =1cm
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Saprobic on leafy substrata. Sexua l state: Ascomata
(242-)303–340μm high×(218-) 238–263(−331) μm diam.,
superficial, solitary, sometimes clustered, gregarious,
subglobose, oval to obovoid, with soft texture, dark brown,
pale brown above, with a central papillate ostiole, with rare
setae. Setae 61–80μm long, dark brown, tapering towards
subacute apex, widest at base, septate. Peridium 39–44μm
wide, composed of brown to reddish-brown cells of textura
angularis. Hamathecium comprising numerous, 1–2μm wide,
filiform, septate, branched, hyaline pseudoparaphyses. Asci
(90-)96–135×15–20μm (
x¼ 112 19μm , n=20), 8-spored,
bitunicate, elongate to cylindric-clavate, thickened at apex,
with (7-)20–31.5μm long pedicels, ocular chamber not ob-
served. Ascospores 42–61×5–8.5μm (
x¼ 53 6μm , n=20),
2–3-seriate, elongate-fusiform, subcylindrical, tapering to-
wards subacute ends, (8-)9–10-septate, not constricted at sep-
tum, hyaline, smooth-walled. Asexual state: hyphomycetous,
helicosporous. Conidiophores macronematous, thick-walled,
septate, branched, brown to moderately brown. Conidiogenous
cells monoblastic to polyblastic, terminal or intercalary, den-
tate, brown to dark brown, darkened at basal septum. Conidia
(17-)22–28μm diam., conidial filaments 4–5μm wide
(
x¼ 24 5μm , n=20), helicoid, tightly coiled 2–3½ times,
hyaline to pale brown, tapering toward flattened end and scar,
multiseptate, not constricted at septum, smooth-walled.
Cultural characteristics (based on dried culture): Colonies
dark brown, superficial and partly immersed in mycelium,
with dark brown, septate, branched hyphae.
Material examined: BERMUDA, Devonshire Marsh,
Devonshire Parish, on leaves of Sabal bermudiana L.H. Bai-
ley (Arecaceae), coll. R.P. Korf and A.Y. Rossman (AR-BER
12), on 18 January 1980; det. A.Y. Rossman (BPI 1104599,
dried culture: BER 12 80–4).
Notes: Helicoma rugosa based on Tubeufia rugosa C.
Booth was originally described as Sphaeria helicoma W.
Phillips & Plowr. (1877). Pirozynski (1972) redescribed and
transferred the fungus to Tubeufia based on that helicosporium
asexual morph. Subsequently, Crane et al. (1998)transferredit
to Thaxteriella because of the pigmented ascomata and
peridial wall structure. In addition, Thaxteriella helicoma also
produced a helicoid asexual state in culture (Fig. 15). Accord-
ing to molecular analysis (Clade A, Fig. 2), the p utative
isolates of Thaxteriella helicoma (JCM 2739 and UBC
F13877) cluster in Helicoma sensu stricto, while the strain
from UBC F13877 clustered with the generic type species
Helicoma muelleri CBS 964.69 with strong support (100 %
BS and 1.00 PP). The placement of Helicoma rugosa in
Helicoma is well-supported by morphology and molecular
phylogeny. Although Sphaeria helicoma is an older name it
cannot be used to represent this species as it would result in a
tautonym; the name of a genus and species that are the same
are not allowed in botanical nomenclature. Therefore we used
the next available synonym which is Helicoma rugosa.
Helicoma miscanthi (W.H. Hsieh, Chi Y. Chen & Sivan.),
Boonmee & K.D. Hyde, comb. n ov., Index Fungorum
numbeter: I F 55063 7, Faces offunginumber : FoF 00180
Fig. 16
≡ Tubeufia miscanthi W.H. Hsieh, C.Y. Chen & Sivan.,
Mycol. Res. 102: 234 (1978)
Saprobic
on dead wood. Sexual state: Ascomata 285–
358(− 373) μ m high ×299– 339(− 386) μ m diam.
(
x¼ 330 334μm ), superficial, solitary, scattered, globose-
subglobose to ovate, flat to slightly shrunken on apex, lacking
obvious setae, hyphae developing from ascomatal base on sub-
strate, brown to dark brown when fresh, with a waxy covering on
ascomal wall, pale brown when dry, darkened at centre, ostiolate.
Peridium 51–58μm wide, comprising several layers of brown to
red brown cells of textura angularis. Hamathecium comprising
numerous, ca.3–4μm wide, long filiform, septate, unbranched,
hyaline pseudoparaphyses. As ci (142-)154–188(−194)×
19–26(−30) μm (
x¼ 172 24μm , n=20), 8-spored, bitunicate,
cylindric-clavate, rounded at apex, mostly apedicellate, ocular
chamber not observed. Ascospor es (122-)131–165(−187.5)×
7–10μm (
x¼ 147 9μm , n=20), fasciculate, cylindric-clavate,
elongate, tapering towards rounded ends, slightly curved, 19–
23(−25)-septate, slightly constricted, hyaline, with minutely glo-
bose cells, smooth-walled. Asexual state: Unknown.
Cultural characteristics: Ascospores germinating on MEA
within 12 h and ge rm tubes produced from both ends.
Colonies growing on MEA slowly, less than 5 mm in 7 days
at 28 C, slightly effuse, undulate to fimbriate at margin, dark,
with white or pale brown aerial mycelium, asexual spores not
formed within 60 days.
Material examined: THAILAND, Chiang Mai, Muang,
Huai Kok Ma, Doi Su Thep, N18°48.365′ E98°54.522′,elev.
1015 msl., on dead wood of an unidentified tree, 21 April 201 1,
Saranyaphat Boonmee, DST-03 (MFLU 11–0134, epitype
designated here; PDD 104453, iso-epitype), ex-type living
culture = MFLUCC 11–0375 = BCC 52033 = ICMP 20072.
Notes: One fresh tubeufia-like specimen with globose,
light-coloured ascomata, numerous pseudoparaphyses,
cylindrical, elongated asci and cylindrical, many septate,
hyaline ascospores is identical with Tubeufia mis c an t hi
(Hsieh et al. 1998). Tubeufia miscanthi was transferred
to Taphrophila (T. miscanthi) by Réblová and Barr
(2000) as it has elongate fusiform and fasciculate asco-
spores arranged in the ascus. However, Tubeufia
miscanthi is morphologically different from the type
species Taphrophila cornu-capreoli (Scheuer 1988;
Hsieh et al. 1998). O ur data indi c at es that Taphrophila
miscanthi should be referred to Helicoma as it lies in
Clade A, supported with 95 % BS and 1.00 PP (Fig. 2 ).
Samuels et al. (1979), and Barr (1980) considered
T. miscanthi to be identical to T. paludosa but t hi s is
not supported here. We epitypify this name here with
the new collection from Thailand.
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Helicoma siamense Boonmee & K.D. Hyde, sp. nov.,
MycoBank: MB 804557, Facesoffunginumber: FoF 00181
Figs. 17–18
Etymology: in reference to its occurrence in Thailand (Siam).
Holotype:MFLU10–0053
Saprobic on dead wood. Sexual state: Ascomata 221–
306μm high×180–268μm diam. (
x¼ 270 233μm ), superfi-
cial, solitary, scattered, with basal subiculum, globose to
subglobose, reddish-brown to dark brown, with a central ostiole,
coveredbybrownmycelium,collapsingwhendry.Peridium
30–48μm wide, composed of several-layers of cells of textura
angularis, with outer layer cells darkened and inner layer cells
pale brown to hyaline. Hamathecium comprising numerous,
wide, filiform, septate, branched, hyaline pseudoparaphyses.
Asci (123-)127.5– 139(− 144)×(14-)17– 20μ m
(
x¼ 133:5 18μm , n=20), 8-spored, bitunicate, cylindrical
to clavate, short-pedicellate,with a rounded apex. Ascospores
(49.5-)59–64.5(−68)×5–7.5μm (
x¼ 62 6:5μm , n=20),
Fig. 16 Helicoma miscanthi (MFLU 11–0134, epitype). a Herbarium
specimen. b Ascomata on substrate. c L.S. of ascoma. d Peridium. e
Pseudoparaphyses. f, g Asci. h, i Ascospores. j Germinating ascospore. k,
l Colonies on MEA from surface and reverse. Scale bars: b–c =100μm,
d =40μm, e =5μm, f–j =50μm, k–l =10mm
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Fig. 17 Helicoma siamense (MFLU 10–0053, holotype). a Ascomata (arrows). b L.S. of ascoma. c Peridium. d Pseudoparaphyses embedded in a
gelatinous matrix. e–g Asci. h–n Ascospores with multiple cells. Scale bars: a–b =100μm, c, h–n =20μm, d =5μm, e–g =50μm
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overlapping 2–3-seriate, cylindric-fusiform, tapering towards
rounded ends, widest at supramedian part, slightly curved, 11 –
12-septate, not constricted at septa, hyaline, smooth-walled.
Asexual state: hyphomycetous, helicosporous. Conidiophores
macronematous, 2–3μm wide, erect, smooth, pale brown to
brown, smooth. Conidiogenous cells holoblastic. Conidia
helicosporous, 24–36μm diam. when coiled, filaments 7–
10μm wide, coiled 2–3 times, multiseptate, slightly constricted,
darkened at septa, partly heavily pigmented, verrucose, rough-
walled, brown to reddish-brown, narrow at base and wide at
upper half on uncoiling.
Cultural characteristics: Ascospores germinating on MEA
within 24–36 h and germ tubes produced from many cells.
Colonies growing on MEA slowly, 6 mm in 7 days at 28 C,
flat, sparsely hairy , fimbriate, dark brown. Mycelium partly
superficial, partly immersed, composed of branched, septate,
smooth, pale brown to reddish-brown, red pigmented hyphal
filaments.
Material examined: THAILAND, Chiang Rai, Muang,
Khun Korn Waterfall, elev. 671 msl., N19°51–54′ E
99°35.39′, on d ecaying wood of an unidentified tree, 13
November 2009, Saranyaphat Boonmee, KK-09 (MFLU
Fig. 18 Helicoma siamense (MFLU 10–0053, holotype). a Germinating ascospore. b, c Colonies on MEA, surface and reverse. Note black colonies. d
Mycelium development in culture. e–l Conidiophores and conidia. Scale bars: a, g–l =20μm, b–c = 10 mm, d–f =5μm
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10–0053, holotype; PDD 104454, isotype); ex-type living
culture = MFLUCC 10–0120 = BCC 52298 = IFRD 2184.
Notes: This taxon is morphologically similar to Tubeufia
and was initially identified as a species of Tubeufia sensu
stricto (Samuels et al. 1979; Sivanesan 1984;Rossman
1987). Helicoma siamense is different from T. javanica in size,
shape and dimensions of the ascomata and ascospores and
produces helicoma-like conidia that are multiseptate, helicoid
with coloured, verrucose, conidial filaments. Phylogenetic
analysis shows that H. siamense clusters with Helicoma sensu
stricto (Clade A, Fig. 2) with high support (97 % BS and 1.00
PP). It differs morphologically and phylogenetically from other
taxa in this Clade and thus is introduced as a new species.
Helicoma chiangraiense Boonmee & K.D. Hyde, sp.
nov., MycoBank: MB 804558, Facesoffunginumber:FoF
00182 Fig. 19
Etymology: In reference to its occurrence in Chiangrai,
Thailand.
Holotype:MFLU10–0048
Saprobic on dead wood . Sexual state: Ascomata
(183-)240– 285 μ m high ×(189-)241– 296 μ m diam.
(
x¼ 242:5 254μm ), superficial, solitary, scattered,
subglobose, ellipsoidal-ovate, with few hyphae developing
from ascomatal base on substrate, dark brown to black, vel-
vety, ostiolate. Peridium 28–34μm wide, comprising 3–4
layers, composed of cells of textura angularis, with inner cells
brown and outer cells dark brown. Hamathecium comprising
ca.0.5– 1 μ m wide, numerous, filiform, hyaline
pseudoparaphyses. Asci (146-)153–179.5×18–23(−26) μm
(
x¼ 163:5 21μm , n=20), 8-spored, bitunicate, cylindrical,
apically thickened and rounded, with a 14.5–24μm long ped-
icel. Ascospores (63-)72–93.5×5–7.5μm (
x¼ 81 6μm , n=
20), overlapping fasciculate, cylindric-fusiform, with tapering
and rounded ends, straight to slightly curved, 11–13-septate,
not constricted at septa, hyaline, smooth-walled. Asexual
state: Unknown.
Cultural characteristics: Ascospores germinating on MEA
within 12 h and germ tubes produced at both ends. Colonies
growing slowly on MEA, reaching less than 5 mm in 7 days at
28 C, effuse, velvety, with a convex surface, with dark con-
tinuous margin, asexual spores not formed within 60 days.
Material examined: THAILAND, Chiang Rai, Muang, Doi
Pui, elev. ca.403–936 msl., on dead wood of Fagaceae,15
August 2009, Saranyaphat Boonmee , DP-01 (MFLU 10–
0048 holotype; PDD 104455 isotype); ex-type living culture
=MFLUCC10–0115 = BCC 39624, 39625 = IFRD 2195 =
ICMP 20068.
Notes: Helicoma chiangraiense differs from other species
of Helicoma in having a sexual morph of dark brown to black,
shiny, non setose ascomata with distinct ostioles (Fig. 19a).
Helicoma chiangrai ense groups with the Helicoma sensu
stricto (Clade A, Fig. 2) with high support (92 % BS and
0.98 PP).
Helicoma fagacearum Boonmee & K.D. Hyde,sp.nov.,
MycoBank: MB 804557, Facesoffunginumber: FoF 00183
Fig. 20
Etymology: in reference to its occurrence on Fagus.
Holotype:MFLU11–0138
Saprobic on dead wood. Sexual state: Ascomata 256–
300(− 324) μ m high ×235– 258(− 265) μ m diam.
(
x¼ 282 251μm ), superficial, s olitary, scattered, with a
subiculum composed of dark brown hyphae, with black stalk,
globose to subglobose, broadly ellipsoid-ovate, yellowish or
brownish, pale brown, ostiolate, collapsing, brown when dry.
Peridium 43–51μm wide, comprising several layers of angu-
lar and prismatic cells, with inner cells pale brown, with outer
cells reddish-brown to brown. Hamathecium with numerous,
ca.1.5μm wide, filiform, septate, branched, hyaline
pseudoparaphyses. Asci (121-)134–160(−179)×10–14μm
(
x¼ 144 12μm , n=20), 8-spored, bitunicate, elongate-cy-
lindrical, apically thickened when young, pedicellate, rounded
at apex. Ascospor es (63-)71–86.5×4–6.5μm (
x¼ 78 5μm ,
n=20), overlapping 2–3-seriate, cylindric-fusiform, tapering
towards rounded ends, straight to slightly curved, 9–12(−13)-
septate, not constricted at septum, hyaline, pale brown at
maturity, guttulate when young, smooth-walled. Asexual
state: Unknown.
Cultural characteristics: Ascospores germinating on MEA
within 12 h and germ tubes produced at both ends. Colonies
growing on MEA slowly, reaching less than 5 mm in 7 days at
28 C, slightly convex on surface, with undulating margin,
white or pale brown to dark brown, turning dark with age,
asexual spores not formed within 60 days.
Material examined: THAILAND, Chiang Rai, Muang, Doi
Pui, elev. ca.403–936 msl., on dead wood of Fagaceae,10
May 2011, Saranyaphat Boonmee, DP-04 (MFLU 11–0138
holotype; PDD 104456 isotype); ex-type living culture =
MFLUCC 11–0379 = BCC 52030 = ICMP 20071.
Notes: Helicoma fagacearum was found on wood of
Fagaceae in northern Thailand. Morphologically, this species
is similar to Helicoma miscanthi in its coloured ascomata with
yellowish white surface, brown peridium and pale brown
ascospores. An asexual morph did not develop in culture.
Our phylogenetic study shows that Helicoma fagacearum
groups with Helicoma sensu stricto (Clade A, Fig. 2)witha
good support (89 % BS and 0.99 PP).
Helicomyces Link, Mag. Gesell. naturf. Freunde, Berlin
3(1–2): 21 (1809), Facesoffunginumber: FoF 00212
Saprobic on woody substrata. Mycelium composed of part-
ly immersed and partly superficial, pale brown, septate,
sparsely branched hyphae, with masses of crowded conidia.
Sexual state: Ascomata superficial, seated on a thin
subiculum, uniloculate, subglobose, clavate to obovate, cori-
aceous, solitary to gregarious, pale brown to reddish-brown,
centrally ostiolate, perhaps collapsing when dry, with radiat-
ing mycelium or appendages at base. Peridium composed of
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cells of textura angularis, with external pale yellow, or brown-
ish to brown layer, with inner hyaline layer. Hamathecium
comprising numerous 1–2 μm wide, filiform septate,
branched, anastomosing, hyaline pseudoparaphyses. Asci 8-
spored, bitunicate, fissitunicate, saccate or cylindric-clavate,
apically rounded, with or without an ocular chamber, long
pedicellate. Ascospores overlapping fasciculate, elongate,
Fig. 20 Helicoma fagacearum (MFLU 11–0138, holoty pe). a
Ascomata on substrate. b L.S. of ascoma. c Peridium. d
Pseudoparaphyses. e–g Asci. h– j Ascospores. k Germinating
ascospore. l, m Colonies on MEA from surface and reverse. Asexual
state not produced. Scale bars: a–b =100μm, c =40μm, d =5μm, e–g =
50μm, h–k =20μm, l–m =10mm
Fig. 19 Helicoma chiangraiense (MFLU10–0048, holotype). a Super-
ficial ascomata on substrate. White oozing mass of ascospores at apex of
ascomata. b L.S. of ascoma. c. C.S. of peridium. d Pseudoparaphyses
embedded in a mucilaginous matrix. e, f Asci. g– i Ascospores. j
Germinating ascospore. k, l Colonies on MEA from surface and reverse,
colonies dark brown. Scale bars: a–b, e–f =100μm, c =20μm, d =5μm,
g–j =50μm, k–l =10mm
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cylindric-subfusiform or narrowly oblong, tapering towards
narrow, subacute ends, 10–11-septate, not constricted at sep-
tum, hyaline, occasionally pale brown, guttulate when imma-
ture, smooth-walled. Asexual state: Conidiophores
macronematous, erect, short, septate, thick-walled, pale to
moderately dark brown. Conidiogenous cells monoblastic or
polyblastic, terminal or intercalary, integrated, subhyaline,
denticulate. Conidia loosely coiled 2½–4 times, becoming
loosely uncoiled in water, conidial filament 4.5–6μm wide,
rounded at apical end, hyaline to pale yellow, multiseptate,
with up to 56-septate, slightly constricted at septum, smooth-
walled.
Type species: Helicomyces roseus Link
Notes: The genus Helicomyces Link (1809) typified by
H. roseus Link is characterized by slender, loosely coiled,
multiseptate, hyaline, helicoid conidia that are filamentous
and white to pinkish in mass. Helicomyces roseus is
frequently a ssociated with an asexual state referred to as
Tubeufia cylindrothecia as first determined by Seaver and
Waterston (1940) and later by Barr (1980). Tsui and Berbee
(2006) and Tsui et al. (2006, 2007) have shown that species
referred to as Helicomyces are polyphyletic with sexual states
in Acanthostigma and Tubeufia. According to our phy-
logenetic study (Fig. 2 ), we therefore treat Clade G as
Helicomyces sensu stricto based on authentic sequences
of Helicomyces roseus CBS 283.51 with 76 % BS and
0.99 PP support. Two isolates of Tubeufia paludosa
(CBS 120503 as epitype and AR 4206) and two species
of Helicosporium (H. indicum an d H. talbotii)cluster
with an isolate of H. roseus CBS 283.51 in Helicomyces
sensu stricto.
Type species: Helico myces rose us Link, Mag. Gesell.
naturf. Freunde, Berlin 3(1– 2): 21 (1809),
Facesoffunginumber: FoF 00184 Fig. 21
Saprobic on woody substrata.
Mycelium partly immersed,
partly superficial, pale brown, septate, sparsely branched hy-
phae, with masses of crowded conidia. Sexual state: Un-
known. Asexual state: Conidiophores erect, 3–4×2–3μm
(
x¼ 4 3μm , n=5), septate, thick-walled, pale to moderately
dark brown, macronematous, short. Conidiogenous cells
monoblastic or polyblastic, terminal or intercalary, integrated,
subhyaline, denticulate. Conidia (40-)47–65(−75) μm diam.,
and conidial filament 4.5–6μm wide (
x¼ 54 6μm , n=20),
with (298.5-)360–398μm long, loosely coiled 2½–4times,
becoming loosely uncoiled in water, rounded at apical end,
bright, hyaline to pale yellow, multiseptate, up to 56-septate,
slightly constricted at septum, smooth-walled.
Material examined: USA, North Carolina, Bearwallow
Cove, on rotten wood, W.W. Diehl, 4 January 1927, Detr:
J.A. Stevenson (BPI 421361).
Helicomyces paludosa (P. Crouan & H. Crouan),
Boonmee & K.D. Hyde, comb. nov., Index Fungorum
number: IF 550689, Facesoffunginumber: FoF 00185 Fig. 22
≡ Nectria paludosa P. Crouan & H. Crouan, Florule du
Finistere p. 38. 1867
≡ Tubeufia paludosa (P. Crouan & Crouan) Rossman,
Mycologia 69(2): 383 (1977)
Saprobic on decorticated or decaying woody and herba-
ceous substrates, in terrestrial habitats, widespread in temper-
ate to tropical regions. Sexual state: Ascomata (267-)278–
297(−312) μm high×180–203μm diam., superficial, seated
on a thin subiculum, uniloculate, subglobose, clavate to ob-
ovate, coriaceous, solitary to gregarious, pale brown to red-
dish-brown, centrally ostiolate, collapsing when dry, with
radiating mycelium or appendages at base. Peridium com-
posed of cells of textura angularis, with outer pale yellow,
or brownish to brown layer, with inner hyaline layer connect-
ed with membranous tissues. Hamathecium comprising nu-
merous, 1–2μm wide, filiform, septate, branched, anastomos-
ing, hyaline pseudoparaphyses. As ci 112–152(−158)×12–
16μm (
x¼ 128 14μm , n =20), 8-spored, bitunicate,
fissitunicate, saccate or cylindric-clavate, sessile or with a
pedicel, sometimes as long as ca.21–24(−33) μm,apically
rounded, with or without an ocular chamber. Ascospores
(53-)59–73(−78)×4–6μm (
x¼ 65 5μm , n=20), overlap-
ping fasciculate, elongate, cylindric-subfusiform or narrowly
oblong, tapering towards n arrow, subacute ends, (9-)10–
11(−13)-septate, not constricted at median septum, hyaline,
occasionally pale brown, guttulate when immature, smooth-
walled. Asexual state: Unknown.
Material examined: USA, Tennessee, Blount Co., Great
Smoky Mountains National Park, 25 mi W Gatlinburg, Cades
Cove, Gregory Ridge Trail, alt. 1950 ft. 35°33′45″N83°50′
45″W, on rotten wood of unindentified angisoperm, A.Y.
Rossman, 6 September 2005 (BPI 871087), living culture =
CBS 120503 = AR 4206.
Notes: Nectria paludosa P Crouan & H Crouan was placed
in Tubeufia as T. paludosa (P. Crouan & H. Crouan) Rossman
based on its bitunicate asci (Rossman 1977). Tubeufia
paludosa was also considered an earlier name for T. javanica,
by Rossman (1977). Barr (1980)consideredT. paludosa dis-
similar from T. javanica, but accepted the synonymy in view
of the similar coloured ascomata and cylindrical ascospores.
The original descriptions of T. paludosa and T. javanica are
similar in having ovoid, translucent, light-colored ascomata,
filiform, cylindrical asci and filiform, hyaline and guttulate
ascospores (Saccardo 1883; Saccardo and Sydow 1899).
However, the illustrations of Tubeufia javanica in Penzig
and Saccardo (1904)andT. paludosa in Samuels et al.
(1979)andSivanesan(1984) indicate there are differences in
structure of the ascomata. Samuels et al. (1979)examinedand
described T. paludosa, and considered the asexual state found
near the ascomata to be helicomyces-like. Tsui et al. (2006,
2007), Tsui and Berbee (2006), Schoch et al. (2009) and
Promputtha and Miller (2010) investigated the molecular phy-
logeny of Tubeufia paludosa. Their phylogenetic analyses
274
Fungal Diversity (2014) 68:239–298
suggest that T. paludosa is related with the helicosporous
asexual morphs. We therefore consider the collection BPI
871087 to be Tubeufia paludosa (CBS 120503 as epitype).
On the basis of phylogenetic placement (Clade G, Fig. 2), two
cultures of Tubeufia paludosa (from CBS 120503 and AR
4206) are related with an asexual species Helicosporium
indicum Rao & D. Rao with 73 % BS and 0.96 PP supports.
An authentic strain of Helicomyces roseus (CBS 283.51) and
Helicosporium talbotii (MUCL 33010) cluster with 92 % BS
support and also form a subclade with Tubeufia paludosa with
76 % BS and 0.99 PP support within Helicomyces sensu
stricto (Clade G).
Helicosporium Nees, Syst. Pilze (Würzburg): 68 (1816)
[1816–17], Facesoffunginumber: FoF00213
Helicotrichum Nees & T. Nees, Nova Acta Acad. Caes.
Leop.-Carol. 9: 246 (1818)
?Drepanospora Berk. & M.A. Curtis [as ‘Drepanispora’],
in Berkeley, Grevillea 3(no. 27): 105 (1875)
Saprobic on leaves, bark, and twigs, or on stromata of
Diatrype stigma, Diatrypaceae (Xylariales), on decorticated
or decaying wood, in terrestrial habitats or submerged in
freshwater, widespread in temperate to tropical regions. Sex-
ual state: Ascomata superficial, solitary, scattered, globose-
subglobose, bright yellow brown to yellow orange, collapsing
when dry, darkened near ostiole. Peridium of several layers,
composed of bright yellow cells of textura prismatica to
angularis. Hamathecium comprising numerous, filiform, sep-
tate, branched, hyaline pseudoparaphyses. Asci 8-spored,
bitunicate, cylindric-clavate, apedicellate, thickened at apex,
with an acute ocular chamber. Ascospores biseriate, elongate-
fusiform, tapering towards narrowly subacute ends, trans-sep-
tate, hyaline. Asexual state: hyphomycetous, helicosporous.
Fig. 21 Helicomyces roseus
(BPI 421361). a Conidia arise
directly from hyphal cells on
natural substrata. b, c
Conidiophores with attached
conidia. d–g Conidia. Scale bars:
a, g =100μm, b–f =50μm
275
Fungal Diversity (2014) 68:239–298
Conidiophores macronematous, mononematous, setiferous,
erect, septate, unbranched, dark brown, fertile in middle,
sterile, tapering towards narrow subacute at apex, smooth-
walled, arising directly on substrate, from thick-walled, close-
ly septate, recumbent hyphae, crowded or in fascicles, glis-
tening, light coloured. Conidiogenous cells polyblastic, inter-
calary, rarely terminal, with lateral minute denticles each with
single conidium. Conidia coiled 3½–4 ½ times, tightly to
loosely coiled, rounded at apical end, truncate at base, hyaline,
7–13-septate, slightly constricted at septa, smooth-walled.
Type species: Helicosporium vegetum Nees
Notes: Helicosporium was introduced by Nees (1817)
based on H. vegetum. At present Helicosporium species are
polyphyletic within the family Tubeufiaceae and are related
with sexual states in the genera Acanthostigma,
Acanthostigmina and Tubeufia (Tsui and Berbee 2006; Tsui
et al. 2006,Fig.2). In our analysis, the type speci es of
Helicosporium, H. vegetum is linked with Tubeufia cerea
(Berk. & M.A. Curtis) Höhn. (Clade D, Fig. 2)withhigh
support (99 % BS and 1.00 PP) and thus synonymized. In
Helicosporium ascomata are bright yellow brown to yellow
orange and formed on stromata of other fungi, and ascospores
are elongate-fusiform, tapering towards narrowly subacute
ends, trans-septate, hyaline, with hyaline, helicoid conidia
and borne directly on small denticles on setiferous conidio-
phores which are terminally sterile. The ex-type cultures of
Helicoma vaccinii (CBS 216.90) and Acanthostigma
patagonica (MVB 573 BBB) cluster in Clade D (Fig. 2)with
strains of Helicosporium sensu stricto. Helicoma vaccinii
produces terminal conidia (Carris 1989) while no asexual state
is known for Acanthostigma patagonica (Sánchez et al. 2012).
Type species: Helicosporium vegetum Nees, Syst. Pilze
(Würzburg): 68 (18 16) [18 16–17], Facesoffunginumber:
FoF 00186 Figs. 23–24
≡ Tubeufia cerea (Berk. & M.A. Curtis) Höhn., Sber. Akad.
Wiss. Wien, Math.-naturw. Kl., Abt. 1 128: 562 (1919)
=Sphaeriacerea
Berk. & M.A. Curtis, Grevillea 4(no.
31): 108 (1876)
Saprobic on leaves, bark, and twigs, or on stromata of
Diatrype stigma, Diatrypaceae (Xylariales), on decorticated
or decaying wood, in terrestrial habitats, or submerged in
freshwater, widespread in temperate to tropical regions. Sex-
ual state: Ascomata 180–211μm high×186–226μm diam.,
superficial, solitary, scattered, globose-subglobose, bright yel-
low brown to yellow orange, collapsing when dry, darkened
near ostiole. Peridium 23–26μm wide, comprising several
layers, composed of bright yellow cells of textura prismatica
to angularis. Hamathecium comprising numerous, 1–1.5μm
wide, filiform, septate, branched, hyaline pseudoparaphyses.
Asci 68–82(−92)×(7-)8–12μm (
x¼ 75 10μm , n=20), 8-
spored, bitunicate, cylindric-clavate, apedicellate, thickened
at apex, with an acute ocular chamber. Ascospores 35–46×
3–5μm (
x¼ 39 4μm , n=10), biseriate, elongate-fusiform,
tapering towards narrowly subacute ends, 8–9-septate, slightly
Fig. 22 Helicomyces paludosa
(BPI 871087, CBS 120503; re-
illustrated from Hyde et al. 2013).
a Ascomata seated on sparse hy-
phae. b Squash mount of ascoma
showing peridium. c
Pseudoparaphyses. d Asci with
thickened apex. e–g Ascospores.
Note asci and ascospores in cot-
ton blue reagent. Scale bars:
a =100μm, b, e–g =20μm,
c =5μm, d =50μm
276
Fungal Diversity (2014) 68:239–298
curved, hyaline, smooth-walled. Asexual state: hyphomy-
cetous, h elicosporo us. Conidiophores (98-)107–220μm
long×3–5μm (
x¼ 148 4μm , n =20), macr onematous,
mononematous, setiferous, erect, septate, unbranched, dark
brown, fertile in middle, sterile, tapering toward narrow sub-
acute at apex, smooth-walled, arising directly on substrate,
from thick-walled, closely septate, repentent hyphae, crowded
or in fascicles, glistening, light coloured. Conidiogenous cells
polyblastic, intercalary, rarely terminal, with lateral minute
denticles each with single conidium. Conidia 10–15μm diam.
and conidial filament 1–2μm wide (
x¼ 13 1:5μm , n=20),
coiled 3½–4½ times, tightly to loosely coiled, rounded at
apical end, truncate at base, 7–13-septate, slightly constricted
at septa, hyaline, smooth-walled.
Notes:VonHöhnel(1919) transferred Sphaeria cerea
Berk. & M.A. Curtis to Tubeufia as T. cerea, based on
bitunicate asci, and narrow elongate, multi-septate ascospores.
Tubeufia cerea is often found on stromata of species belong-
ing in Diatrypaceae (Xylariales). Morphologically, Tubeufia
cerea was distinguished from all other species in the genus by
its yellow brown to yellow orange ascomata (Bigelow and
Barr 1963; Booth 1964; Samuels et al. 1979;Barr1980).
Booth (1964) provided a description and illustration of
Tubeufia cerea based on the type specimen and provided a
list of synonyms. The asexual state found near the ascomata
were identified as Helicosporium vegetum Nees by
Booth (1964), Samuels et al. (1979)andBarr(1980).
Phylogenetic analysis links Tubeufia cerea with
Helicosporium ve getum with high support (Tsui and
Berbee 2006;Tsuietal.2006) and this is supported
in our study (Clade D, Fig. 2). Tubeufia cerea (Berk. &
M.A. Curtis) Höhn. is therefore treated as a synonym of
Helicosporium vegetum.
Material examined: USA, Virginia, Falls Church, on rotten
wood, C.L. Shear, May 1936; Detr: J.A. Stevenson (BPI
447464, material of asexual state); Virginia, Shenandoah Na-
tional Park, Beams Gap, on stromata of Diatrype stigma
(Diatrypaceae, Xylariales), on dead wood, A.Y. Rossman
(1925), 19 June 1983; Detr: A.Y. Rossman (BPI 1107327,
sexual state of Tubeufia cerea).
Kamalomyces R.K. Verma, N. Sharma & Soni, Forest
Fungi of Central India: 196 (2008)
Fig. 23 Helicosporium cereum
(BPI 1 107327 as Tubeufia cerea).
a Ascomata on fungal stromata. b
Section of ascoma. c Peridium. d
Pseudoparaphyses. e, f Asci. g–i
Ascospores. Scale bars: a =
200μm, b =100μm, c =50μm,
d =5μm, e–i =20μm
277
Fungal Diversity (2014) 68:239–298
Saprobic on dead bamboo. Sexual state: Ascomata
forming on a subiculum of crowded black mycelium,
superficial, clustered t o solitary, globose to subglobose,
stalked, ostiolate. Peridium comprising 3–4 layers of
darkened cells of textura angularis,dullatmargin.
Hamathecium comprising filiform, branched
pseudoparaph yses, embedded in a g elatinous matrix.
Asci 8-spored, bitunicate, thick-walled, saccate-clavate,
pedicellate, with an ocular chamber. Ascospores fusi-
form to clavate, slightly tapering toward rounded ends,
trans-septate with crowded septa, straight or slightly
curved, upper part broad, hyaline (from Boonmee
et al. 2011). Asexual state: Unknown.
Notes: This genus has ascomata that form on a subiculum
of crowded, black mycelium and fusiform to clavate, trans-
septate, hyaline ascospores with crowded septa (Boonmee
et al. 2011). There are presently two species (Dubey and
Neelima 2013), both from bamboo.
Neoacanthostigma Boonmee, Bhat & K.D. Hyde, gen.
nov., Index Fungorum number: IF 550576,
Facesoffunginumber: FoF00214
Etymology: neo – a new genus similar to Acanthostigma.
Saprobic on dead wood. Sexual state: Ascomata superfi-
cial, solitary, scattered, globose to subglobose, reddish-brown
to dark brown, to black, surrounded by black shining setae,
tapering towards an acute tip, ostiolate. Peridium composed of
several layers of brown cells of textura angularis.
Hamathecium comprising numerous, filiform, septate,
branched, hyaline pseudoparaphyses. Asci 8-spored,
bitunicate, cylindric-clavate, with a short rounded pedicel,
with thick and rounded apex, ocular chamber not observed.
Ascospores fasciculate, cylindrical, narrowly fusiform, taper-
ing towards narrow, subacute ends, trans-septate, hyaline,
smooth-walled. Asexual state: hyphomycetous,
helicosporous. Conidiophores micronematous, holoblastic,
polyblastic, dentate on creeping hyphae, up to 5μm long,
Fig. 24 Helicosporium vegetum
(BPI 447464) a Conidiophores
with conidia on natural substrate.
b, c Conidiophores with minute
denticles (arrows). d–f Conidia
stained in cotton blue in f. Scale
bars: a =100μm, b =50μm,
c =5μm, d–f =10μm
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Fungal Diversity (2014) 68:239–298
hyaline. Conidia helicosporous, coiled in 1–2½ dimensional
times, tapering towards rounded ends, multiseptate, hyaline,
with mucilaginous pads at ends.
Type species: Neocanthostigma fusiforme Boonmee, Bhat
&K.D.Hyde
Notes: The type species of Neoacanthostigma, N. fusiforme,
clusters with the type strain of Acanthostigma
septoconstrictum Promp. & A.N. Mill. in an individual clade
in the multigene analysis (Clade M, Fig. 2)withhighsupport
(99 % BS and 1.00 PP). These taxa appear to be related,
therefore, we synonymize Acanthostigma septoconstrictum
under Neoacanthostigma based on phylogenetic suggestion.
Two strains of Acanthostigma filiforme (ANM101, ex-type
and ANM514) also cluster in Clade M, although with weak
support (Fig. 2) and is synonymised under Neoacanthostigma.
Neoacanthostigma is similar to Acanthostigma in that
ascomata are covered with setae, asci are bitunicate, asco-
spores are cylindrical to narrowly fusiform and the asexual
state is helicosporous (Réblová and Barr 2000). Species in this
genus would previously have been referred to Acanthostigma,
while their asexual states are helicoma-like or helicomyces-
like. These genera are morphologically distinct. In
Neoacanthostigma theascomatahavelongsetaeandthe
ascospores are narrowly fusiform to symmetrical, while in
Acanthostigma the ascomatal setae are short and the asco-
spores are asymmetrical, fusiform to clavate (Réblová and
Barr 2000; Boonmee et al. 2011). The hyaline conidiophores
are moniliaceous and micronematous formed directly from
mycelium, not typical of Helicoma, Helico myces or
Helicosporium. Phylogenetically species of Neocanthostigma
in Clade M are basal to Clades A-L (Fig. 2).
Neoacanthostigma fusiforme Boonmee, Bh at & K.D.
Hyde, sp. nov., Index Fungorium numb er: IF 5505 77,
Facesoffunginumber: FoF 00187 Figs. 25–26
Etymology: in reference to the symmetrical fusiform
ascospores.
Holotype:MFLU11–1146
Saprobic on dead wood . Sexual state: Ascomata
(111-)126– 138 μ m
high × 98– 100(− 125) μ m diam.
(
x¼ 125 107μm ), superficial, solitary, scattered, globose
to subglobose, reddish-brown to dark brown to black,
surrounded by shining black setae (27-)48–73μm long, taper-
ing towards an acute tip, ostiolate. Peridium 13–14μm wide,
composed of several layers of brown cells of text ura
angularis. Hamathecium comprising ca. 1.5–2μm wide, nu-
merous, filiform, septate, branched, hyaline
pseudoparaphyses. Asci 71– 84 ×10– 11 (− 12) μ m
(
x¼ 79 11μm , n=20), 8-spored, bitunicate, cylindric-cla-
vate, with a short rounded pedicel, with thick and rounded
apex, ocular chamber not observed. Ascospores (32-)40–48×
3–4.5μm (
x¼ 43 4μm , n=20), fasciculate, cylindrical, nar-
rowly fus iform, tapering towards narrow, subacute ends,
straight to slightly curved, 5–7-se ptate, n ot constricted at
septum, with mucilaginous pads at ends, hyaline, smooth-
walled. Asexual state : hyphomycetous, helicosporous.
Conidiophores up to 5μm long, micronematous, holoblastic,
polyblastic, dentate on creeping hyphae, hyaline, smooth-
walled. Conidia helicosporous, (17-)23–30μm diam. when
coiled, conidial filaments 2μm wide, coiled in 1–2½ dimen-
sional times, tapering toward rounded ends, multiseptate, not
constricted at septa, hyaline, smooth-walled.
Cultural characteristics: Ascospores germinating on MEA
within 8 h and germ tubes produced at both ends. Colonies
growing on MEA slowly, less than 5 mm diam in 7 days at 28
C, slightly raised-radially with lobate to entire edge, grayish to
pale brown, laterally becoming dark brown. Mycelium devel-
oping on substrate, superficial, with hyaline to pale brown
hyphae.
Material examined: THAILAND, Chiang Mai, Mae
Taeng, Mushroom Research Centre, N19°17.123′ E98°44.
009′,elev.ca.900msl.,ondeadwoodofanunidentifiedtree,
23 June 2011, Saranyaphat Boonmee, MRC-03(T) (MFLU
11–1146 holotype; PDD 104457 isotype); ex-type living
culture = MFLUCC 11–0510 = BCC 52383 = ICMP 20073.
New combinations:
Neoacanthostigma septoconstrictum (Promp. & A.N.
Mill.) S. Boonmee & K.D. Hyde, comb. nov., Index
Fungorum number:IF550683
≡ Acanthostigma septoconstrictum Promp. & A.N. Mill.,
Mycologia 102(3): 579 (2010)
Neoacanthostigma filiforme (Promp. & A.N. Mill.) S.
Boonmee & K.D. Hyde, comb. n ov., Index Fungorum
number IF 550684
≡ Acanthostigma filiforme Promp. & A.N. Mill.,
Mycologia 102(3): 575 (2010)
Podonectria Petch, Trans. Br. mycol. Soc. 7: 146 (1921)
Associated with scale insects.
Sexual state: Ascomata su-
perficial, scattered, solitary, subglobose-globose, light yellow-
reddish, covered with light brown mycelium. Peridium com-
posed of several-layers of pale yellowish brown cells of
textura angularis. Hamathecium comprising filiform, septate,
branched, hyaline pseudoparaphyses, embedded in a gelati-
nous matrix. Asci 8-spored, bitunicate, cylindro-clavate, short
pedicellate. Ascospores 2–3-seriate, clavate to fusiform, hya-
line to pale brown, trans-septate, wall minutely verruculose
(from Boonmee et al. 2011). Asexual state: parasitic on scale
insectis, hyphomycetous, phragmosporous, Tetracrium,by
forming holoblastic conidia and borne on sympodially prolif-
erating conidiogenous cells (from Pirozynski 1977).
Notes: The genus is tentatively included in T ubeufiaceae
pending fresh collections and phylogenetic analysis. Its sexual
state is compatible with T ubeufiaceae, however, the asexual
states are atypical in producing conidia on sympodial conid-
iophores (Pirozynski 1977; Sivanesan 1984; Rossman 1987).
Type species: Podonectria coccicola (Ellis & Everh.)
Petch, Trans. Br. mycol. Soc. 7: 146 (1921)
279
Fungal Diversity (2014) 68:239–298
Tamhinispora K.C. Rajeshk. & Rahul Sharma,
Mycosphere 4(2): 166 (2013), Facesoffunginumber:
FoF00215
Saprobic on submerged wood. Mycelium mostly semi-
immersed or immersed, single or interwoven. Stroma none.
Setae and hyphopodia absent. Sexual st ate: Unk nown.
Fig. 25 Neocanthostigma fusiforme (MFLU 11–1146, holotype). a
Ascomata on substrate (arrows). b L.S. of ascoma. c Setae. d Peridium.
e Pseudoparaphyses. f–h Asci. i–k Ascospores. Note spores becoming
yellow when stained in Melzer’s reagent in Fig. k. Note the mucilaginous
pads at the ends. Scale bars: a–b =100μm, c, f–h =50μm, d, i–k =
20μm, e =5μm
280
Fungal Diversity (2014) 68:239–298
Asexual state: Conidiophores absent. Conidiogenous cells,
semi-macronematous, unbranched, intercalary in hyphae, al-
most sessile. Conidia solitary, dry, simple, mostly ovoid or
irregular, dictyoseptate, smooth-walled, mature conidia dark
brown to blackish with apical appendages. Apical appendages
rudimentary or well-developed, arising from apex of conidia,
diverging or radiating, pale to dark brown, septate, apex
rounded (from Rajeshkumar and Sharma 2013).
Type species: Tamhinispora indica K.C. Rajeshk. & Rahul
Sharma
Notes: This unique genus with only an asexual state was
described from submerged wood in India (Rajeshkumar and
Sharma 2013). The dark brown conidia are adorned with
brown to blackish apical appendages, albeit small, are remi-
niscent of asco mata of Tubeufiaceae. In the phylogenetic
analyses it groups basal to Clade F and Clade G of Tubeufia
and Helicomyces, with 0.99 PP support clade (Fig. 2).
Type species: Tamhinispora indica K.C. Rajeshk. & Rahul
Sharma, Mycosphere 4(2): 167 (2013), Facesoffunginumber:
FoF 00188 Fig. 27
Thaxteriella Petr., Annls m ycol. 22(1/2): 63 ( 1924),
Facesoffunginumber: FoF00216
Saprobic on dead bark. Mycelium forming a dense velvety
subiculum on surface of bark, comprising black hyphae. Sex-
ual state: Ascomata superficial or slightly embedded on a
basal subiculum, solitary or densely gregarious, globose to
turbinate, black, shiny, often with small apical papillae at
central ostiolate apex, often collapses when dry. Peridium
comprising two strata, outer layer of thick-walled, dark brown
cells of textura angularis and inner layer of brown cells of
textura globosa. The apical region is composed of less thick-
walled cells that disintegrate to form an ostiole. Hamathecium
composed of filamentous, septate, branched, anastomosing,
hyaline pseudoparaphyses. Asci 8-spored, bitunicate, cylindri-
cal to broadly clavate or subclavate, usually with a short
pedicel, thickened at apex with an ocular chamber. Ascospores
cylindrical to long fusiform, tapering towards rounded to sub-
acute ends, trans-septate, hyaline. Asexual state: hyphomy-
cetous, helicosporous. Conidiophores erect, up to 180μm
long, brown to dark brown, light-coloured at apex. Conidia
17–21μm diam., pleurogenous, developing on cylindr ical
pegs arising form cells of conidiophores, 1½–1¾ times coiled,
5–12-septate, hyaline to pale olivaceous brown (from
Sivanesan 1984).
Notes: No sequences exist in GenBank for the type species
of Thaxteriella, T. corticola and other species in the genus.
Thaxteriella pezizula was included in this genus by Petrak
(1953) based on morphological consistency with the genus, as
recognized by globose or oblate globose, darkly pigmented,
shiny, ascomata, collapsing in the middle when dry and fili-
form ascospores (Berkeley 1876;Linder1929;Barr1980;
Sivanesan 1984). In addition, Linder (1929) determined
Fig. 26 Neocanthostigma
fusiforme (MFLU 11–1146,
holotype). a Germinating
ascospore. b Colonies on MEA.
Note colonies are grayish to pale
brown. c, d Growth of asexual
state on plant tissues produced on
the media with and without
substrate. e Aerial mycelium in
culture. f Conidiophores formed
on hyphae (arrows). g–j Conidia.
Scale bars: a =20μm, b–d =
10 mm, e–f =5μm, g–j =10μm
281
Fungal Diversity (2014) 68:239–298
that Thaxteriella pezizula produced a helicoma-like hy-
phomycetous asexual state. Thaxteriella may therefore
be a synonym of Helicoma. Boonmee et al. (2011)
introduced Thaxteriella inthanonensis as a new species
in Thaxteri ella, and this species clusters with Helicoma
sensu stricto as reported
Type species: Thaxteriella corticola Petr., Annls mycol.
22(1/2): 63 (1924)
Thaxteriella pezizula (Be rk. & M.A. Curtis) Petr.,
Sydowia 7(1–4): 110 (1953), Facesoffunginumber:FoF
00189 Fig. 28
≡ Sphaeria pezizula Berk. & M.A. Curtis, Grevillea 4(no.
31): 106 (1876)
Saprobic on woody substrates.Myceliumforming a dense
velvety subiculum on the surface of the bark, comprising
black hyphae. Sexual state: Ascomata 236–374×365–
422μm diam., superficial, seated on mycelium, solitary to
clustered, scattered on substrate, globose to subglobose, tur-
binate, dark brown to black, shiny, ostiolate, collapsing on
drying. Peridium 44.5–55μm, several layers, somewhat thick-
ened, composed of dark brown cells of textura angularis.
Hamathecium of 2–3.5μm wide, filiform, septate, branched,
hyaline pseudoparaphyses. Asci (102-)113–146(−151)×25–
34μm (
x¼ 128 30μm , n=20), 8-spored, bitunicate, broadly
cylindric-subclavate, shorted pedicellate, thickened at apex,
lacking a distinct ocular chamber. Ascospores 47–61×9.5–
13μm (
x¼ 53 11μm , n=20), overlapping 2–3-seriate, cy-
lindrical to long fusiform, tapering towards rounded ends, 7-
septate, straight or slightly curved, not constricted at septa,
hyaline to pale-yellowish. Asexual state: hyphomycetous,
helicosporous. Conidiophores erect, up to 180μm long, brown
to dark brown, light-coloured at apex. Conidia 17–21μm
diam., pleurogenous, develop on cylindrical pegs arising from
cells of conidiophores, 1½–1¾ times colied, 5–12-septate,
hyaline to pale olivaceous brown (from Sivanesan 1984).
Material examined: Unknown location, on dead wood of
undetermined timber, collector and date collected are un-
known (BPI 800530).
Thaxteriellopsis Sivan., Panwar & S.J. Kaur, Kavaka 4: 39
(1977) [1976]
Saprobic on dead wood. Sexual state: Ascomata superfi-
cial, borne on a thin, dark brown subiculum, solitary or
scattered, globose to subglobose, reddish-brown to dark
brown, with brown to dark brown, septate, setae, mostly at
the apex. Peridium comprising 3–4 layers of red brown to
dark br own cells of textura an gularis. Hamathecium
comprising filiform, hyaline pseudoparaphyses, embed-
ded in a gelatinous matrix. Asci 8-spored, bitunicate,
cylindrical to clavate, apically rounded, short pedicel-
late, with long apical region with amorphous contents.
Ascospores 2–3-ser iate, fus iform to c lavate, broader
above, straight to slightly curved, 5-septate, constricted
Fig. 27 Tamhinispora indica (AMH 9555: holotype; re-illustrated from Rajeshkumar and Sharma 2013). a–c Conidial spores formed on natural
substrate. d–h Immature and mature conidia with developing apical appendages. Scale bars: e–i =20μm, j =10mm
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at septum, hyaline, smooth-walled. Asexual state:hy-
phomycetous, helicosporous. Micronematous conidia-
like structures, develop directly from hyphae in culture
(from Boonmee et al. 2011).
Notes: Thaxteriellopsis is a strongly-supported genus in the
phylogenetic analysis with 100 % BS and 1.00 PP (Clade B,
Fig. 2). The type species, T. lignicola, is associated with a
moorella-like asexual morph (Subramanian and Sekar 1982).
Boonmee et al. (2011) found that Thaxteriellopsis lignicola
produced micronematous, brown, helicosporous, septate,
conidia-like structures directly on the hyphae in culture. For
an account of this genus, see Boonmee et al. (2011).
Type species: Thaxteriellopsis lignicola Sivan., Panwar &
S.J. Kaur, Kavaka 4: 39 (1977) [1976]
Genera excluded from Tubeufiaceae (Tubeufiales)
Paranectriellaceae S. Boonmee & K.D. Hyde
This new family was introduced by Hyde et al. (2013)to
accommodate two genera, namely Paranectriella and
Puttemansia.
Paranectriella (Henn. ex Sacc & D. Sacc) Höhn., Sber.
Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119: 899 (1910)
Notes: The genus Paranectriella based on P. j u ru a n a was
first recognized as a subgenus of Paranectria by Hennings
(1904), which was validated by Sacca rdo and Saccardo
(1905). Von Höhnel (1910) later raised Paranectriella to
generic rank. Pirozynski (1977) recognized Paranectriella
and redescribed the type species, distinguishing
Paranectriella having bitunicate asci from Paranectria with
untunicate asci. Barr (1980) suggested that Paranectriella be
included in T ubeufiaceae due to its habit as a tropical hyper-
parasite. Rossman (1987) accepted the genus Paranectriella
in Tubeufiaceae,describingP. juruana as a parasitic species
Fig. 28 Thaxteriella pezizula
(BPI 800530). a, b Ascomata
collapsing when dry (in a). c L.S.
of ascoma. d Peridium. e
Pseudoparaphyses. f, g Asci. h–j
Ascospores. Scale bars: a–c =
200μm, d =50μm, e =5μm,
f–g =50μm, h–j =20μm
283
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with bright white to yellow ascostromata containing many
locules in a peripheral layer, bitunicate asci,
pseudoparaphyses and ascospores with polar spine-like ap-
pendages. The ascostromata occur on hyaline, relatively loose
mycelium appearing intermediate between an ascostromata
and a subiculum. The associated asexual states of
Paranectriella are hyphomycetous, staurosporous, referred
to as Araneomyces and Titaea (Saccardo 1876; von Höhnel
1909b; Kirschner and Piepenbring 2006;Hydeetal.2011).
The description and illustration here are based on an authentic
specimen of Paranectriella juruana (BPI 632134). Morpho-
logically, P. juruana is characterized by bright ascostromata
and ascospores with appendaged ends. These characters are
atypical of genera in Tubeufiaceae, which have cylindrical
ascospores and hyphomycetous, helicosporous asexual states.
Paranectriella was placed in a new family Paranectriellaceae
in Hyde et al. (2013).
Type species: Paranectriella juruana (Henn.) Henn. ex
Piroz., Kew Bull 31: 598 (1977)
≡ Paranectria juruana Henn., Hedwigia 43(4): 245 (1904)
Notes: AdetaileddescriptionofParanectriellaceae based
on P. juruana, was given in Hyde et al. (2013).
Puttemansia Henn., Hedwigia 41: 112 (1902),
Facesoffunginumber: FoF00217
Parasitic on lower surface of leaves of Nectandra sp.
Sexual state: Ascostromata up to 1 mm diam., superficial,
solitary, sometimes in groups, pale yellow to vinaceous buff,
stipitate, covered by hairy setae, white to hyaline, septate,
multi-loculate. Locules globose-subglobose, slightly flattened
at apex, non-ostiolate. Peridium composed of cells of textura
prismatica to oblonga, hyaline to pale brown. Hamathecium
comprising numerous filiform, branched, anastomosing, hya-
line pseudoparaphyses, embedded in a gelatinous matrix. Asci
8-spored, bitunicate, oblong, elongate to cylindrical-clavate,
rounded at apex, obtuse at base. Ascospores 1–2-seriate, fusi-
form, widest near central septum, tapering towards narrow
ends, 3-septate, slightly constricted at septum, with continu-
ous basal appendage, hyaline, smooth-walled. Asexual state:
hyphomycetous, staurosporous, Guelichia, Tetranacrium
(Hyde et al. 2011).
Type species
: Puttemansia lanosa Henn.
Notes:Hennings(1902) introduced the genus Puttemansia
as a member of Pezizaceae. Clements and Shear (1931), and
Rogerson (19 70)placedPuttemansia in Hypocreaceae,
(Hypocreales) based on charactistics of hairy appendages
and bright ascomata. Pirozynski (1977) and Rossman (1978)
provided detailed descriptions of several species of
Puttemansia and found that all species had bitunicate asci.
Rossman (1978) suggested that Puttemansia was similar to
Podonectria and Tubeufia in Tubeufiaceae having bright,
fleshy ascomata, bitunicate asci and multi-septate ascospores
(Barr 1980). Rossman (1987) provided detailed descriptions
of six sp ecies and accepted them in Tubeufiacea e, with
Puttemansia lanosa Henn. was treated as a synonym of
Puttemansia albolanata (Speg.) Höhn. The asexual states of
Puttemansia are considered to be the hyphomycetous genera
Guelichia Speg., Tetranacrium H.J. Huds. & B. Sutton and
Titaea Sacc., as they were commo nly associated with
Puttemansia lanosa (Hennings 1902; Petrak and Sydow
1936; Barr 1980; Rossman 1987; Hyde et al. 2011). The
description and illustration herein, is from an authentic spec-
imen of Puttemansia lanosa (BPI 632856). Morphological
features of the fungus are cup-shaped ascostromata, white
hairy m ycelium, and broadly fusiform appendaged a sco-
spores, characters that are atypical of T ubeufiaceae.
Puttemansia lanosa is similar to Paranectriella juruana in
that ascostromata are light coloured and ascospores three
septate with appendages. However, they differ as
ascostromata of Puttemansia lanosa contain fewer locules
(2–10) and ascospores have a basal spine-like appendage. In
Paranectriella juruana the ascostromata contain many locules
(ca. 30), while ascospores have spine-like appendages at both
ends. The genus Puttemansia therefore, is referred to the
Paranectriellaceae, based on morphological similarities
(Fig. 90 in Hyde et al. 2013).
Type species: Puttemansia lanosa Henn., Hedwigia 41:
112 (1902), Facesoffunginumber: FoF 00190 Fig. 29
= Guelichia paradoxa Speg., Anal. So c. cient. argent.
22(2): 174 (1886)
Parasitic on lower surface of leaves of Nectandra sp.
Sexual state: Ascostromata up to 1 mm diam., superficial,
solitary, sometimes grouped, pale yellow to vinaceous buff,
stipitate, covered by flexuous setae, 200–386×6–9μm,white
to hyaline, septate, multi-loculate. Locules (244-)259–
306(− 567) μ m high×(314-)371– 421μ m diam.
(
x¼ 334 370μm ), globose-subglobose, slightly flattened
at apex, non-ostiolate. Peridium 64–73μm wide, composed
of cells of textura prismatica to oblonga, hyaline to pale
brown. Hamathecium comprising numerous filiform, 1–2μm
wide, branched, anastomosing, hyaline pseudoparaphyses,
embedded in a gelatinous matrix. Asci (104-)112–
133(−140) ×16.5– 22μm (
x¼ 120 19μm ), 8-spored,
bitunicate, oblong, elongate to cylindrical-clavate, rounded
at apex, obtuse at base. Asc ospores (30-)38–50(−56)×9–
11μm (
x¼ 42 10μm ), 1–2-seriate, fusiform, widest near
central septum, tapering towards narrow ends, 3-septate,
slightly constricted at septum, with continuous basal append-
age, hyaline, smooth-walled. Asexual state: hyphomycetous,
staurosporous, Guelichia, Tetranac rium (Hyde et al. 2011).
Material examined: COSTA RICA, San Pedro de San
Ramon, on leaves of Nectandra sp.Rol.exRottb.
(Lauraceae), 8 October 1926, Alberto M. Brenes, Fungi
costaricensis 157 (BPI 632856).
Pleosporales incertae sedis
Based on Hyde et al. (2013)thePleosporales comprises 31
families whose members are saprobic on dead plant material
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in freshwater, marine, or terrestrial environments, and a large
number, especially as asexual states, are pathogens on living
plants (Zhang et al. 2009; Boonmee et al. 2012; Liu et al.
2012). Characteristics include perithecial ascomata usually
with distinct ostioles, abundant pseudoparaphyses, pedicel-
late, cylindro-clavate asci usually with distinct ocular cham-
bers and various ascospore types. Glaxoa and Rebentischia
are not typical of any family in Pleosporales and are therefore
placed in Pleosporales incertae sedis.
Glaxoa Cannon, Syst. Ascom. 15(1–2): 122 (1997)
Notes: Glaxoa is a monotypic genus, typified by Glaxoa
pellucida (Cannon 1997). Herein, we provide a drawing from
Cannon (1997), because the type material of G. pellucida,
which we have examined (IMI 362099), is in poor condition.
Fig. 29 Puttemansia albolanata
(BPI 632856) a, b Ascostromata
on leaf. c Section of ascoma. d
Loose mycelium of ascostromata.
e Peridium. f Pseudoparaphyses.
g–i Immature and mature asci. j–
m Ascospores with basal spine-
like appendage. Scale bars: a =
5mm,b =500μm, c =200μm, d–
e =100μm, g–i =50μm, f =5μm,
j–m =20μm
285
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Glaxoa pellucida can clearly be distinguished from all genera
in Tubeufiaceae by its 1-septate ascospores. Cannon (1997)
placed Glaxoa in Tubeufiace ae because it was similar to
Letendraea, which is presently treated as a member of
Pleosporales (Boonmee et al. 2011). Glaxoa is therefore
placed in Pleosporales incertae cedis.
Type species: Glaxoa pellucida P.F. Cannon, Syst. Ascom.
15(1–2): 122 (1997), Facesoffunginumber: FoF 00191 Fig. 30
Rebentischia P. Karst., Fungi Fenniae Exsiccati, Fasc. 9:
no. 881 (1869), Facesoffunginumber: FoF00218
Saprobic on decaying wood. Sexual state: Ascomata su-
perficial, solitary or scattered, globose-subglobose, black, co-
riaceous, ostiolate, sometimes collapsed when dry. Peridium
thick, composed of cells of textura angularis,withdarkbrown
to black cells. Pseudoparaphyses numerous, 1–2μm wide,
flexuous, hyaline, septate, branched. Asci 8-spored, bitunicate,
fissitunicate, cylindric-clavate, short pedicellate, apically
thickened. Ascospores biseriate, elongate-obovoid, trans-sep-
tate, brownish to dark brown, light at apex, base tapering to
hyaline appendage. Asexual state: Unknown.
Type species: Rebentischia pomiformis P. Karst.
Notes: Karsten (1869) introduced the genus Rebentischia
based on R. pomiformis. Rebentischia has been placed in
different families (Saccardo 1877, 1883;Müller1950;Von
Arx and Müller 1975). Barr (1980)includedRebentischia in
T ubeufiaceae based on its temperate saprobic habit, darkly
pigmented ascomata, bitunicate asci, and elongated, multi-
septate ascospores. In addition, Barr (1980)placed
Rebentischia pomiformis
in synonymy with R. massalongi
as first suggested by Von Arx and Müller (1975). Only two
accepted species, R. massalongi (Mont.) Sacc.
(=R. pomiformis P. Karst.) and R. unicaudata (Berk. &
Broome) Sacc. were discussed by Barr (1980). Barr (1980)
also suggested that coelomycetous taxa, such as asteromella-
like fungi found near the ascomata might be the asexual state
of Rebentischia massalongi. Rebentischia pomiformis has
darkened ascospores with a setiform basal appendage appears
typical of Pleosporales. The genus is therefore included in
Pleosporales incertae sedis.
Type species: Rebentischia pomiformis P. Karst., Fungi
Fenniae Exsiccati, Fasc. 9: no. 881 (1869),
Facesoffunginumber: FoF 00192 Fig. 31
Saprobic on decaying wood of Acer platanoides. Sexual
state: Ascomata 249–287μm high×270–296(−323) μm
diam., superficial, solitary or scattered, globose-subglobose,
bluish-black, velvety, coriaceous, ostiolate, sometimes col-
lapsed when dry. Peridium thick, 25–28μm wide, composed
of dark brown to black-cells cells of textura angularis .
Hamathecium comprising numerous, 1–2μm wide, flexuous,
hyaline, septate, branched pseudoparaphyses. Asci (99-)106–
126 ×15–20μm (
x¼ 112:5 17μm , n =10), 8-spored,
bitunicate, fissitunicate, cylindric-clavate, short pedicellate,
apically thickened, with an ocular chamber. Ascospores 24–
29(−35)×6–10μm (
x¼ 27 8μm , n=20), biseriate, elongate-
obovoid, 4-septate, slightly constricted at septum, brownish to
dark brown, lighter at apex, base tapering to 4–8.5(−14) μm
long, narrow, hyaline appendage, smooth-walled. Asexual
state: Unknown.
Material examined: FINLAND, Abo, on wood of Acer
platanoides L. (Sapindaceae), 7 April 1861, P.A. Karsten,
Fig. 30 Glaxoa pullucida (a,b
IMI362099, holotype, c–e
redrawn from Cannon 1997). a
Herbarium packet. b Surface
view. c Ascoma. d Asci and
interascal tissue. e Ascospores.
Scale bars: a–b =detailedof
Glaxoa pullucida specimen, c =
50μm, d–e =10μm
286
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Finland Fungi 881, Farlow Herbarium, Harvard University,
Herbarium of F. Theissen (FH, holotype).
Dothideomycetes genera incertae sedis
One-hundred and sixteen genera are listed as incertae sedis
in the class Dothideomycetes in Lumbsch and Huhnd orf
(2010). Below we add a further four genera previously placed
in Tubeufiaceae that cannot be placed in any family or order
with certainty.
Acanthostigmella Höhn., Annls mycol. 3(4): 327 (1905),
Facesoffunginumber: FoF00219
Saprobic on dead stems of grasses. Sexual state: Ascomata
superficial, solitary, scattered, globose-subglobose, dark
brown, with 23–74μm long dark brown, apically acute,
septate setae, darkened papilla and ostiole. Peridium
thin-walled, composed of brown to red brown, 3–
6.5μ m diam. cells of textura angularis. Hamathecium
lacking pseudoparaphyses. Asci 8-spored, bitunicate, ob-
long, subclavate to broadly obovoid, with a blunt, wide,
rounded pedicel, almost a pedicellate. Ascosp ores 2–3-
seriate in ascus, ellipsoid, fusoid, ends rounded, 2–3-
Fig. 31 Rebentischia pomiformis (FH, holotype). a Scattered ascomata on host. b Ascoma in section. c Peridium. d Wide flexuous pseudoparaphyses.
e–g Immature and mature asci. h–l Ascospores with basal appendage. Scale bars: a =200μm, b =100μm, c, e–g =50μm, d =5μm, h–l =10μm
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septate, constricted at septum, yellow brown, smooth-
walled. Asexual state: see notes.
Type species: Acanthostigmella genuflexa Höhn.
Notes: Acanthostigmella was proposed by Von Höhnel
(1905) based on the type species A. genuflexa.Materialfrom
FH was not in good condition and devoid of any fungi. M.E.
Barr had annotated the specimen as containing broken
fragments bearing a few perithecia and small spored species
of Gaeumannomyces. Two slides were present in the exsiccata
and Barr (1977) obtained her description from these. A more
recent collection (IMI 252801) similar to the species drawn
from the FH type slides by Barr (1977) is illustrated below.
Morphologically, this specimen is different from all genera in
T ubeufiaceae with small ascomata containing a distinct papil-
la surrounded by setae, lack of pseudoparaphyses, broadly
obovoid to fusoid asci, and 2–3-septate, yellow brown asco-
spores (von Höhnel 1905; Barr 1977). Barr (1977, 1980)and
Rossman (1987) suggested that A. genuflexa would be better
accommodated in Herpotrichiellaceae (Chaetothryriales)
rather than Tubeufiaceae. Based on molecular data of
Acanthostigmella brevispina M.E. Barr & Rogerson,
Untereiner et al. (1995) placed the taxon in Dothideomycetes
(Figs. 22 and 23 in Untereiner et al. 1995). Whether this
species is related to A. genuflexa is debatable because of the
presence of pseudoparaphyses, a hyphomycetous asexual
state and fungicolous habitat. The placement of this genus
therefore has to be Dothideomycetes genera incertae sedis.
The asexual states of Acanthostigmella are listed as hypho-
mycetous, helicosporous, dictyosporous and xenosporium-
like (Hyde et al. 2011). However, no asexual state is linked
to the generic type.
Type species: Acanthostigmella genuflexa Höhn., Annls
mycol. 3(4): 327 (1905), Facesoffunginumber: FoF 00193
Fig. 32
≡ Acanthostigma genuflexum (Höhn.) Sacc. & Trotter, in
Saccardo, Syll. fung. (Abellini) 22: 209 (1913)
≡ Tu beufia genuflexa (Höhn.) Arx & E. Müll., Stud.
Mycol. 9: 83 (1975)
Saprobic on dead stems of grasses. Sexual state: Ascomata
20–79μm high×3 6–77μm diam ., superficial, solitary,
scattered, globose-subglobose, dark brown; with surface se-
tae, papillate and with darkened, 13–20μm diam. ostiola, with
setae 23–74μm long, dark brown, apical acute, septate.
Peridium thin-walled, ca.4.5–5μm wide, composed of brown
to reddish-brown cells 3–6.5μm diam. of textura angularis.
Hamathecium lacking pseudoparaphyses. Asci 19–23×8–
11μm (
x¼ 21 9μm , n=10), 8-spored, bitunicate, oblong,
subclavate to broadly obovoid, with a blunt, wide, rounded
pedicel, almost apedicellate. Ascospores 7–11 ×2–3μm
(
x¼ 9 3μm , n=10), 2–3-seriate in ascus, ellipsoid,
fusoid, rounded at both ends, 2–3-septate, constricted
at septum, yellow brown, smooth-walled. Asexual state:
Unknown.
Material examined: AUSTRIA, Langenschönbichl, Tulln,
3 June 1905, v Höhnel (FH, holotype,poorspecimen);UK,
Suffolk, Halesworth (Canal side), on dead stem of Phalaris
arundinacea L. (Poaceae), M.B. and J.P. Ellis, 21 June 1979
(IMI 252801).
Chaetocrea Syd., Annls mycol. 25(1/2): 18 (1927),
Facesoffunginumber: FoF00220
Fungicolous associated with black areas of fungi on leaves.
Sexual state: Ascomata superficial, solitary or gregarious,
globose-subglobose, membranous, white to pale yellow, cov-
ered by apically branched setae. Peridium comprising whitish
yellow cells of textura angularis. Hamathecium comprising
filiform, septate, branched, hyaline pseudoparaphyses, devel-
oping over asci. Asci 8-spored, bitunicate, elongate-clavate,
cylindrical, short pedicellate, apically thickened, without an
obvious ocular chamber. Ascospor es fasciculate, filiform, ta-
pering narrowly towards lower end, with several septa, hya-
line. Asexual state: Unknown.
Type species: Chaetocrea parasitica Syd.
Notes: Chaetocrea was introduced by Sydow (1927)fora
single species Chaetocrea parasitica andplacedinthe
Hypocreaceae as it had bright-coloured ascomata. Chaetocrea
parasitica was described as fungicolous, growing on black-
ened areas of Cyclostomella disciformis which parasitized
leaves of Nectandra sanguine. Rossman et al. (1999)studied
genera in the Hypocreaceae (Hypocreales) and transferred
Chaetocrea to Tubeufiaceae based on its bitunicate asci,
pseudoparaphyses and long ascospores. Chaetocrea
parasitica has predominantly branching, pale yellow setae
atypical of T ubeufiaceae.ThereforeChaetocr ea is treated as
amemberofDothideomycetes
genera incertae sedis.
Type species: Chaetocrea parasitica Syd., Annls mycol.
25(1/2): 19 (1927), Facesoffunginumber: FoF 00194 Fig. 33
Fungicolous associated with black areas of Cyclostomella
disciformis on leaves of Nectandra sanguine. Sexual state:
Ascomata 161–187μm high×169–189μm diam., superficial,
solitary or gregarious, globose-subglobose, membranous,
white to pale yellow, translucent or crystal-like, covered by
35–44(−48) μm long×7–10μm wide, apically branched, sep-
tate, pale yellow setae. Peridium 23–26μm wide, 3–4layers
of white-yellow cells of textura angularis. Hamathecium
comprising 2–2.5μm wide, filiform, septate, branched, hya-
line pseudoparaphyses, developing above asci. Asci 95–
108(−121)×17–21μm (
x¼ 102 19μm , n=10), 8-spored,
bitunicate, elongate-clavate, cylindrical, short-pedicellate, api-
cally thickened, without an obvious ocular chamber.
Ascospores (63-)71–83×5–6.5μm (
x¼ 76 6μm , n=10),
fasciculate, filiform, tapering narrowly towards lower end,
Fig. 32 Acanthostigmella genuflexa (a, b holotype FH, c–l Material of
herb. K, as IMI 252801). a, b Herbarium package and specimen. c
Ascomata on substrate. d Close up of ascomata with long setae. e
Peridial wall. f, g Immature and mature asci. h–j Ascospores. Scale
bars: a =500μm, b =100μm, c–d =50μm, e–j =10μm
288
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(6-)7–9-euseptate, not constricted at septum, hyaline, smooth-
walled. Asexual state: Unknown.
Material examined: COSTA RICA, La Caja, near San Jose,
parasitic on Cyclostomella disciformis Pat., growing on leaves
of Nectandra sanguinea Rol. ex Rottb (Lauraceae), 4 January
1925 (No. 166), Acqu. 1978, No. 11007: F. Petrak
Pilzherbarium (W, holotype).
Malacaria Syd., Annls mycol. 28(1/2): 69 (1930),
Facesoffunginumber: FoF00221
Fungicolous associated with Irenina glabra (Meliolaceae);
comprising branched and coloured, superficial mycelium, on
leaves of Coffea robusta, pantropical. Sexual state: Ascomata
superficial, solitary, globose-subglobose, pale brown to or-
ange brown, soft-textured, with a membranous wall, centrally
ostiolate, darkened and collapsing when dry. Peridium com-
posed of 3–5-layers of pale orange cells of textura angularis.
Hamathecium with numerous filiform, septate, unbranched,
hyaline pseudoparaphyses. Asci 8-spored, bitunicate, oblong-
subclavate, rounded at apex, obtuse at base. Ascospores fas-
ciculate, elongate-fusiform, slightly curved, 3–(−4)-septate,
slightly constricted at septa, pale brown, smooth-walled, api-
cally subacute, continuous with basal filiform or thread-like,
hyaline appendage. Asexual state: Unknown.
Type species: Malacaria meliolicola Syd.
Notes: Sydow (1930) introduced the genus Malacaria,
typified by M. meliolicola for species commonly associated
with species of Meliolaceae growing on leaves. Rossman
(1983)treatedM. flagellata as a synonym of M. meliolicola,
based on its similar morphology. She assigned a lectotype for
Paranectria flagellata (≡Malacaria flagellata) and, since the
type of M. meliolicola was lost, she also assigned it as the
neotype of this species ( Rossman 1987). Malacaria
meliolicola was included in Tubeufiaceae based on its soft
ascomata, bitunicate asci and pseudoparaphyses. In this study,
we reexamined the lectoty pe specimen. Malacaria
meliolicola has pale s moke-grey asc ospores and un-
branched sep tate pseu dopara physes and is atypica l of
genera in Tubeufiaceae. The placement of Malacaria is
therefore uncertain and the genus is treated as
Dothideomycetes gen e ra incertae cedis.
Fig. 33 Chaetocrea parasitica
(W, holotype). a Cyclostomella
disciformis on leaf and ascomata
of Chaetocrea parasitica (right
figure) on blackened fungus. b
Apically branching setae. c
Section of ascoma. d Peridium. e
Pseudoparaphyses. f–h Asci. i–k
Ascospores. Scale bars: a =
200μm, b, i–k =20μm, c =
100μm, d, f–h =50μm, e =5μm
290
Fungal Diversity (2014) 68:239–298
Type species: Malacaria meliolicola Syd., Annls mycol.
28(1/2): 69 (1930), Facesoffunginumber: FoF 00195 Fig. 34
= Paranectria flagellata Hansf., Proc. Linn. Soc. London
153(1): 28 (1941)
≡ Malacaria flagellata (Hansf.) Hansf., Mycol. Pap. 15:
128 (1946)
Fungicolous associated with Irenina glabra (Meliolaceae);
comprising branched and coloured, superficial mycelium, on
leaves of Coffea robusta, pantropical. Sexual state: Ascomata
144–170μm high×125–176μm diam., superficial, solitary,
globose-subglobose, pale brown to orange brown, soft in
texture, membranous, centrally ostiolate, darkened and col-
lapsing when dry. Peridium 12–14μm wide, composed of 3–5
layers, of pale orange cells of textura angularis. Hamathecium
of ca. 1–1.5μm wide, numerous, filiform, septate, un-
branched, hyaline pseudoparaphyses. Asci 53–72.5(− 80)×
11–21μm (
x¼ 65 17μm , n=15), 8-spored, bitunicate, ob-
long-subclavate, rounded at apex, obtuse at base. Ascospores
42–51×4–5μm (
x¼ 46 4μm , n=10), fasciculate, elongate-
fusiform, slightly curved, 3–(−4)-septate, slightly constricted
at septum, pale brown, smooth, apically subacute, continuous
with basal ca.43–65μm long, filiform or thread-like, hyaline
appendage. Asexual state: Unknown.
Material examined: UGANDA, Kampala, elev. 4000 ft., on
leaves of Coffea robusta and associated with Irenina glabra
(Berk. & M.A. Curtis) F. Stevens (Meliolaceae), collected by
Hansford G.C., 1871 (K (M): 177970, neotype of Malacaria
meliolicola).
Uredinophila Rossman, Mycol. Pap. 157: 43 (1987),
Facesoffunginumber: FoF00222
Parasitic on Pucciniales on leaves of Dryopteris
tetragona. Sexual state: Ascomata superficial on substrate
and surrounded by rust fungi, developing on a subiculum,
globose to subglobose, solitary, scattered, translucent, yellow
to orange, soft, apex with hyaline hairs. Peridium composed
of hyaline or pale yellow cells of textura angularis.
Hamathecium comprising numerous, 1–2μm wide, cylindri-
cal, branched, hyaline pseudoparaphyses, exending above
asci, embedded in a gelatinous matrix. Asci 8-spored,
bitunicate, cylindrical-clavate, with short knob-like pedicel,
apex rounded. Ascospores fasciculate, helically coiled, fili-
form or fusiform, narrowly elongate, tapering to a narrow
Fig. 34 Malacaria meliolicola
(K (M): 177970, neotype). a
Fungus on black colonies of
Meliolaceae. Orange coloured
mycelium are those of the fungal
parasite. b Section of ascoma. c
Peridium. d Pseudoparaphyses.
e–g Asci. h–k Ascospores with
long, basal appendage. Scale
bars: a =500μm, b =50μm,
c–k =20μm
291
Fungal Diversity (2014) 68:239–298
rounded base, trans-septate, hyaline. Asexual state:
Unknown.
Type species: Uredinophila tropicalis (Speg.) Rossman
Notes:Rossman(1987) introduced the genus Uredinop hila
for two mycoparasitic fungi associated with rusts, i.e. U. erinacea
(Rehm) Rossman a nd U. tropicalis (Speg.) Rossman.
Ur edinophila was excluded from Ophionectria basedonits
bitunicate asci and spirally elongated ascospores (Rossman
1977); and referred to Tubeufiaceae by Rossman (1987)who
provided a key, descriptions and illustrations of the two species.
We reexamined an authentic specimen of U. tropicalis (BPI
632877) listed by Rossman (1987) which has characters that
are atypical of T ubeufiaceae. Therefor e, Uredinoph ila is placed
in Dothideomycetes genera incertae sedis.
Type species: Uredinophila tropicalis (Speg.) Rossman,
Mycol. Pap. 157: 45 (1987), Index Fungorum number:IF
130896, Facesoffunginumber: FoF 00196 Fig. 35
≡ Ophionectria tropicalis Speg., Anal. Soc. cient. argent.
16(5): 242 (1883)
Parasitic associated with Desmella superficialis
(Pucciniales)onleavesofDryopteris tetragona. Sexual state:
Ascomata (122-)134–176(−196) μm high×(111-)141–
167(−196) μm diam. (
x¼ 156 156μm ), superficial on sub-
strate and surrounded by rust fungi, developing on a
subiculum, globose to subglobose, solitary, scattered, translu-
cent, yellow to orange, soft, apex with (16-)21.5–35(−39) μm
long, hyaline hairs, rounded at ends, lacking septa. Peridium
9.5–13μm wide, composed of hyaline or pale yellow, 6–9μm
Fig. 35 Uredinophila tropicalis
(BPI 632877). a Ascomata
associated with a rust. b, c Squash
mount of ascomata. d Close up of
setae and peridial wall. e
Pseudoparaphyses. f, g Asci. h, i
Ascospores. Scale bars: a =
100μm, c, f–i =50μm, d =20μm,
e =5μm
292
Fungal Diversity (2014) 68:239–298
wide, cells of textura angularis. Pseudoparaphyses numer-
ous, 1–2μm wide, cylindrical, cellular, branched, hyaline,
extending above asci, embedded in a gelatinous matrix. Asci
(85-) 92.5–115× 12–15μm (
x¼ 99 13μm ), bitunicate, 8-
spored, cylindrical-clavate, with short knob-like pedicel, with
rounded apex. Ascospores (96-)107.5– 11 3×3 .5– 5μm
(
x¼ 103 4μm ), f asciculate, helically coiled, filiform or
fusiform, narrowly elongate, with rounded apex, tapering to
a narrow rounded base, with 7 or more septa, not constricted at
septa, hyaline, smooth-walled. Asexual state:Unknown.
Material examined: VENEZUELA, Caguita, near Pueto La
Cruz, fungicolous, associated with uredosoris of the fern rust
Desmella superficialis , on leaves of Dryopteris tetragona
(Sw.) U rb. (Dryopteridaceae), H. Sydow, 27 December
1927, Sydow 840 (BPI 632877, issued as Ophionectria
tropicalis).
Ascomycetes genera incertae sedis
Garethjone sia K.D. Hyde, Aust. Syst. Bot. 5(4): 408
(1992)
Notes: This genus was introduced from freshwater by Hyde
(1992), but was later synonymized with Boerlagiomyces by
Stanley and Hyde (1997). The genus is rather distinct and
differs from Boerlagiomyces in having much lighter ascomata,
bicelled asci and large lacunose ascospores. The asci were
described as unitunicate, but this was questioned by Stanley
and Hyde (1997). We re-examined the figures in Hyde (1992)
and the asci appear to be unitunicate (although this cannot be
clearly determined); this is however supported by the fact that
the paraphyses are larg, oval and taper at the ends. We there-
fore treat Gar ethjonesia as a distinct genus which should be
placed in the Ascomycetes, genera incertae sedis.
Boerlagiomyces grandisporus Stanley & K.D. Hyde is also
similar and should probably also be placed in Garethjonesia.
Type species: Garethjonesia lacunosispora K.D. Hyde,
Aust. Syst. Bot. 5(4): 411 (1992)
Doubtful Dothideomycete genera
Amphinectria Speg., Boln Acad. nac. Cienc. Córdoba
26(2–4): 346 (1924)
Notes: We provide above a brief description of
Amphinectria portoricensis based on the protologue of
(1924), the original drawing in the herbarium packet and the
rather depauperate type material. Petrak (1951)examinedthe
type specimen of A. portoricensis, which has immature asci,
and concluded that the species is a lichen. Pirozynski (1977)
later questionably synonymised Amphinectria with
Melioliphila,whileRossman(1987) examined the type spec-
imen of A. portoricensis and could not find any ascomata that
resembled those described by Spegazzini suggesting that the
identity of this species is obscure. Rossman et al. (1999
)
placed A. portoricensis in T ubeufiaceae with uncertainty and
remarked that, until another specimen is located and studied,
Amphinectria will remain an ambiguous member. A second
species in the genus, Amphinectria erubescens,was
transferred to the genus Hydropisphaera (Bionectriaceae),
based on its orange ascomata and unitunicate asci (Vizioli
1923; Rossman et al. 1999). According to the original anno-
tations of Spegazzini (LPS 13394, holotype), this genus and
species is ambiguous and is unlike any other members of
T ubeufiaceae. Therefore we treat Amphinectria as a doubtful
genus.
Type species: Amphinectria portoricensis Speg., Boln
Acad. nac. Cienc. Córdoba 26(2– 4): 346 (1924),
Facesoffunginumber: FoF 00197 Fig. 36
Parasitic on leaf surface of Comocladia glabra (Schult.)
Spreng. Subiculum 5–10 mm diam., orbicular, slightly trans-
parent, inconspicuous, loosely attached to epidermis, with 2–
3μm diam., branching, radial hyphae. Sexual state: Ascomata
200–25 0μ m diam., superficial, globose, solitary, yellow
brown to brown, membranous, developing at centre of
subiculum, minutely papilla, ostiolate, collapsing when dry,
lacking pseudoparaphyses. Asci (70-)75–100×(15-)25–
30μm, 8-spored, bitunicate, cylindrical-ellipsoid, bluntly
rounded at apex, with a short, bifurcate pedicel. Ascospores
(32-)40–45×(6-)8–10μm, 2-seriate in ascus, cylindrical-
fusoid, tapering towards sub-rounded ends, slightly
inequilateral or curved, 7–9-septate, slightly constricted at
septum, hyaline, smooth-walled, lacking a gelatinous sheath
(from Spegazzini 1924). Asexual state: Unknown.
Chaetosphaerulina I. Hino, Bulletin Miyazaki Coll. Agric.
Forest. 10: 62 (1938)
This genus was redescribed and redrawn from the
protologue by Boonmee et al. (2011
). It is similar to several
genera of T ubeufiaceae (e.g. Thaxteriella). The holotype at
YAM appear to have been lost. The genus is therefore treated
as doubtful until it can be rediscovered.
Type species: Chaetosphaerulina yasudai I. Hino, Canad.
J. Plant Sci. 10: 62 (1938)
Melioliphila Speg., Boln Acad. nac. Cienc. Córdoba 26(2–
4): 344 (1924)
Notes: Melioliphila was introduced by Spegazzini (1924)
for mycoparasitic species associated with Meliola
( Meliolaceae). Initially, Melioliphila was placed in the
Hypocreales (Saccardo 1972), but later transferred into
“hypocreoid Dothideales” by Pirozynski (1977). Barr (1980)
transferred Melioliphila to Tubeufiaceae basedonits
mycoparasitic habit and having bitunicate asci and elongate
ascospores. Rossman (1979, 1987) reported that
M. graminicola (F. Stev) Speg. (≡Calonectria graminicola
F. Stev.) is a synonym of M. volutella (Berk. & Broome)
Rossman (≡Calonectria volutella). Melioliphila graminicola
was also placed in T ubeufiaceae by Rossman (1987,1999).
We could not loan the type of M. graminicola due to LPS
policy. Paratype specimens of M. graminicola (BPI632019)
examined were in poor condition. Therefore, we used the
drawings of Spegazzini (1924) and original herbarium packet
(www. cybertruffle.org.uk/spegazzini/ 001626a) to illustrate
293
Fungal Diversity (2014) 68:239–298
M. graminicola. According to Spegazzini (1924),
M. graminicola, has hyaline setae, elliptical-fusoid asci,
fusoid ascospores and lacks pseudoparaphyses, is atypical of
genera in T ubeufiaceae.Therefore,Melioliphila is treated as a
doubtful genus in Dothideomycetes.
Type species: Melioliphila graminicola Speg., Boln Acad.
nac. Cienc. Córdoba 26(2–4): 344 (1924), Facesoffunginumber:
FoF 00198 Fig. 37
Parasitic on Meliola panici (Meliolaceae) growing on
leaves of Lasiacis divaricata in Puerto Rico. Sexual state:
Ascomata 150–200μm diam., globose, developing on a poor-
ly developed, yellowish to fleshy-pink, subiculum, covered by
powder-like, white colonies, with sparsely radiating setae.
Setae 50–100×8–10μm, erect, narrowly tapering towards
apex, hyaline, septate. Hamathecium lacking
pseudoparaphyses. Asci 70–90×10– 14μm, 8-spored,
bitunicate, elliptical-fusoid, short pedicellate, somewhat flat-
tened at apex. Ascospores 30– 40×4– 5 μm, 2-seriate,
narrowly-fusoid, initially with minute appendages at both
ends and appendages disappearing and both ends becoming
blunt when mature, 7–9-guttulate present when immature,
becoming 7–9-septate when mature, hyaline to pale smoky
(from Spegazzini 1924). Asexual state: Unknown.
Notes: The brief description and drawing of Melioliphila
graminicola is provided based on the protologue of
Spegazzini (1924) and the drawing on the herbarium packet
(www. cybertruffle.org.uk/spegazzini/eng/001626a).
Conclusion
Based on molecular phylogenetic studies, family
T ubeufiaceae is placed in order Tubeufiales.Thisrecognition
is strongly supported by morphological characteristics of
asexual and sexual states. The sexual morph of Tubeufiales
Fig. 36 Amphinectria
portoricensis (Redrawn from
LPS 13394, holotype). a, b
Material in LPS packet
(holotype). c, d Substrate and
ascoma. e Asci. f Ascospores
294
Fungal Diversity (2014) 68:239–298
is circumscribed by uniloculate, pseudothecial, superficial
ascomata, which bear setae in some genera, a
pseudoparaphysate hamathecium, bitunicate asci, multi-sep-
tate, hyaline to pale brown cylindrical ascospores and often
connected with hyphomycetous, helicosporous asexual states
(Tsui and Berbee 2006;Tsuietal.2006, 2007; Boonmee et al.
2011).
Furthermore, all genera and new species are provided with
detailed descriptions and illustrations. Phylogenetic placement of
Tubeufiales was determined using combined data set of LSU and
ITS gene regions. Our morpho-molecular analyses confirm that
Tubeufiales, Tubeufiaceae, comprises 19 genera viz.
Acanthohelicospora, Acanthophiobolus, Acanthostigma,
Acanthostigmina, Aquaphila, Boerlagiomyces, Bifrontia,
Chlamydotubeufia , Helicangiospora, Helicoma, Helicomyces,
Helicosporium, Kamalomyces, Neoacanthostigma, Podonectria,
Tamhinispora, Thaxteriella, Thaxteriellopsis and Tubeufia.Pres-
ently, GenBank has over 100 sequences of T ubeufiales including
different gene regions such as LSU, SSU and ITS. In this study
we have used the combined data sets of LSU and ITS gene
regions to resolve genera of Tubeufiaceae. In addition, a fresh
collection of Tubeufia javanica is introduced as the epitype for
Tubeufia sensu stricto.WeexcludeAcanthostigmella,
Amphinectria, Chaetocrea, Chaetosphaerulina, Glaxoa,
Malacaria, Melioliphila, Paranectriella, Puttemansia,
Rebentischia and Ur edinophila from T ubeufiaceae, based on
morphological studies of type species.
Acknowledgments This work was supported by TRF/BIOTEC pro-
gram Biodiversity Research and Training Grant BRT R_251181, BRT
R_253012, USDA-ARS, Systematic Mycology and Microbiology Lab-
oratory (SMML) and partially by the Mushroom Research Foundation,
Chiang Rai Province for financial support. The International Fungal
Research and Development Centre, The Research Institute of Resource
Insects, the Chinese Academy of Forestry (Bailongsi, Kunming 650224,
China) and Mae Fah Luang University (grant for study Dothideomycetes
No. 56101020032) are also thanked for research facilities. Gareth Jones is
supported by the Distinguished Scientist Fellowship Program (DSFP),
King Saud University, Saudi Arabia. K. D. Hyde thanks The Chinese
Academy of Sciences, project number 2013T2S0030, for the award of
Fig. 37 Melioliphila
graminicola (Redrawn from
Spegazzini 1924 and LPS 1626,
www.cybertruffle.org.uk/
spegazzini/eng/001626a). a
Substrate. b, c Ascoma and basal
mycelium. d Setae. e Ascus. f
Ascospores
295
Fungal Diversity (2014) 68:239–298
Visiting Professorship for Senior International Scientists at Kunming
Institute of Botany. Appreciation is extended to the CBS-KNAW Fungal
Biodiversity Centre, The Netherlands, the International Fungal Research
& Development Centre, Research Institute of Resource Insects, Chinese
Academy of Forestry and the State key lab of Mycology, Institute of
Microbiology, Chinese Academic of Science, Beijing, China for provides
molecular sequences. Special thanks also go to Lisa A. Castlebury,
Donald W. Walker, Cesar Herrera, Shanon Dominick, Sajeewa S. N.
Maharachchikumbura, Nalin N. Wijayawardene, Dhanushka Udayanga
and Dimuthu S. Manamgoda for their assistance.
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