Fungal Diversity
DOI 10.1007/s13225-015-0340-y
Towards unraveling relationships
in Xylariomycetidae (Sordariomycetes)
Indunil C. Senanayake 1,2,3 & Sajeewa S. N. Maharachchikumbura 4 & Kevin D. Hyde 1,2,3 &
Jayarama D. Bhat 6,7 & E. B. Gareth Jones 8 & Eric H. C. McKenzie 9 & Dong Qin Dai 1,2,3 &
Dinushani A. Daranagama 3,5 & Monika C. Dayarathne 1,2,3 & Ishani D. Goonasekara 1,2,3 &
Sirinapa Konta 3 & Wen Jing Li 1,2,3 & Qiu Ju Shang 3 & Marc Stadler 14 &
Nalin N. Wijayawardene 3 & Yuan Pin Xiao 3 & Chada Norphanphoun 3 & Qirui Li 3,10 &
Xing Zhong Liu 5 & Ali H. Bahkali 8 & Ji Chuan Kang 10 & Yong Wang 10 & Ting Chi Wen 10 &
Lucile Wendt 15 & Jian Chu Xu 1,2 & Erio Camporesi 11,12,13
Received: 30 May 2015 / Accepted: 29 June 2015
# School of Science 2015
Abstract The classification of subclass Xylariomycetidae
is revisited with additional collections and phylogeny
based on novel rDNA sequence data. Phylogenetic inferences are provided and are based on analysis of 115
sequence data, including new data for 27 strains. An
updated outline to the subclass is presented based on the phylogenies and comprises two orders, 18 families and 222 genera. An account of each order, family and genus in the subclass is given. We accept the orders Amphisphaeriales and
Xylariales based on morphological and phylogenetic evidence. Amphisphaeriales comprises Amphisphaeriaceae,
Bartaliniaceae fam. nov., Clypeosphaeriaceae, Discosiaceae
f a m . n o v. , P e s t a l o t i o p s i d a c e a e f a m . n o v. a n d
Phlogicylindriaceae fam. nov. Xylariales comprises
Apiosporaceae, Cainiaceae, Coniocessiaceae, Diatrypaceae,
Graphostromataceae (doubtful), Hyponectriaceae,
I o d o s p h a e r i a c e a e , L o p a d o s t o m a c e a e f a m . n o v. ,
Melogrammataceae, Pseudomassariaceae fam. nov.,
Vialaeaceae and Xylariaceae. The new genera and species
introduced are Arthrinium hyphopodii, A. subglobosa,
C a i n i a a n t h o x a n t h i s , C i f e r r i a s c o s e a g e n . n o v. ,
C. fluctamurum, C. rectamurum, Discosia neofraxinea,
D. pseudopleurochaeta, Hyalotiella rubi, Seimatosporium
cornii, S. ficeae, S. vitis and Truncatella spartii.
* Kevin D. Hyde
kdhyde3@gmail.com
7
Department of Botany, Goa University, Goa 403 206, India
8
Department of Botany and Microbiology, King Saudi University,
Riyadh, Saudi Arabia
1
Key Laboratory for Plant Biodiversity and Biogeography of East
Asia, Kunming Institute of Botany, Chinese Academy of Science,
Kunming 650201, Yunnan, China
9
Manaaki Whenua Landcare Research, Private Bag
92170 Auckland, New Zealand
10
2
East and Central Asia, World Agroforestry Centre,
Kunming 650201, Yunnan, China
Engineering Research Center for Southwest Bio-Pharmaceutical
Resources, Ministry of Education, Guizhou University,
Guiyang, Guizhou 550025, China
3
Centre of Excellence for Fungal Research, Mae Fah Luang
University, Chiang Rai, Thailand
11
A.M.B. Gruppo Micologico Forlivese, Antonio Cicognani, Via
Roma 18, Forlì, Italy
4
Guizhou Key Laboratory of Agricultural Biotechnology, Guizhou
Academy of Agricultural Sciences, Guiyang 550006, Guizhou,
China
12
A.M.B. Circolo Micologico, Giovanni Carini, 314 Brescia, Italy
13
Società per gli Studi Naturalistici della Romagna,
144 Bagnacavallo, RA, Italy
14
Department of Microbial Drugs, Helmholtz-Zentrum für
Infektionsforschung GmbH, Inhoffenstrasse 7,
38124 Braunschweig, Germany
15
Department of Microbial Drugs, Braunschweig, Helmholtz Centre
for Infection Research (HZI), Braunschweig, Germany
5
6
State Key Laboratory of Mycology,
Institute of Microbiology, Chinese Academy of Sciences,
No 3 1st West Beichen Road, Chaoyang District, Beijing 100101,
China
No. 128/1-J, Azad Housing Society, Curca, P.O. Goa Velha 403108,
India
Fungal Diversity
Keywords Amphisphaeriales . Apiosporaceae .
Bartaliniaceae . Cainiaceae . Ciferriascosea .
Clypeosphaeriaceae . Coniocessiaceae . Discosiaceae .
Diatrypaceae . Hyponectriaceae . Hypoxyloideae .
Iodosphaeriaceae . Lopadostomaceae . Melogrammataceae .
Pestalotiopsidaceae . Phlogicylindriaceae .
Pseudomassariaceae . Vialaeaceae . Xylariaceae .
Xylarioideae
Introduction
The order Amphisphaeriales was introduced by Eriksson and
Hawksworth (1986), and in most subsequent cases was treated
as a synonym of Xylariales. The family Amphisphaeriaceae
was introduced by Winter (1884–1886) as “Amphisphaerieae”,
based on the type genus Amphisphaeria. It was distinguished
from other families in having partially immersed ascomata in the
substrate and from its unique shape of the ostiolar canal.
Amphisphaeriaceae was reintroduced by Müller and von Arx
(1962) based on the presence of a small iodine positive ring or
disc in the ascal apex and in having ascomata immersed under a
clypeus (Kang et al. 1999b). Hawksworth et al. (1995) included
Amphisphaeriaceae in the order Xylariales.
The Xylariales was initially included in the class
Hymenoascomycetes by Barr (1990), but later placed in the
subclass Xylariomycetidae (Eriksson and Winka 1997).
Several of the most well-known species may form conspicuous stromata and are clearly visible on the surface of decaying
plants (Kuhnert et al. 2013). Krug (1978) introduced the family Cainiaceae to accommodate the genus Cainia whose species were previously placed under Amphisphaeriaceae.
During a revision of the genera of Calosphaeriaceae, Barr
et al. (1993) introduced Graphostromataceae as a monotypic
family in the order Xylariales based on the nodulisporium-like
conidial morph and diatrype-like sexual morph having
allantoid ascospores. The family Apiosporaceae was introduced by Hyde et al. (1998). Later, Crous and Groenewald
(2013) used molecular data to confirm the placement of this
family in Xylariales, as a sister group to Amphisphaeriaceae.
Coniocessiaceae was introduced by Asgari and Zare
(2011), placed in Xylariales and comprised of the single genus
Coniocessia with four species. Vialaeaceae was introduced by
Cannon (1995) for species with isthmoid ascospores. Recent
studies have provided phylogenetic evidence that this is a
well-supported family in Xylariales (Shoemaker et al. 2013;
McTaggart et al. 2013; Senanayake et al. 2014).
LSU molecular data have shown a close relationship between Amphisphaeriaceae, Cainiaceae, Clypeosphaeriaceae
and Hyponectriaceae (Jeewon 2001) in Xylariales. Also, in an
attempt to show the evolutionary relationships among the accepted families of this order, Smith et al. (2003) used molecular data, which resulted in showing Xylariales as a
monophyletic group containing Amphisphaeriaceae,
Apiosporaceae, Clypeosphaeriaceae, Diatrypaceae,
Graphostromataceae, Hyponectriaceae and Xylariaceae.
However, the use of 28S and 18S alone proved to be
inadequate for determining the relationships between these
families.
Lumbsch and Huhndorf (2010) treated the order Xylariales
with six families and 154 genera. Currently 11 families have
been listed for Xylariales, namely; Amphisphaeriaceae,
Apiosporaceae, Cainiaceae, Clypeosphaeriaceae,
Coniocessiaceae, Diatrypaceae, Graphostromataceae,
Hyponectriaceae, Melogrammaceae, Vialaeaceae and
Xylariaceae (Maharachchikumbura et al. 2015). Renewed interest in the group Xylariales has resulted in the publication of
monographs for the genera Daldinia, Pestalotiopsis and
Rosellinia (Petrini 2013; Maharachchikumbura et al. 2014;
Stadler et al. 2014). The amphisphaeriaceous members of
Xylariales are however, still relatively poorly understood
and there is no natural classification developed for the group.
As a result of various collections of microfungi from
China, Italy and Thailand, we have encountered several genera of Amphisphaeriales and subjected them to morphomolecular studies. The phylogeny resulting from the analyses
of a combined LSU and ITS gene dataset show that our understanding of the order is fragmentary. In this paper, we resurrect the order Amphisphaeriales and provide a natural classification for the taxa of Xylariomycetidae.
Materials and methods
Specimen collection, morphological examination
and photomicrography
Fresh, plant materials, such as leaves, twigs, stems and bark
were collected from different geographical localities in China,
Italy and Thailand. Fresh materials were placed in paper bags
and all collection details were noted. Specimens were brought
to the laboratory and examined under a stereomicroscope to
observe the characteristics of stromata and fruiting bodies.
Macro-morphological characters were photographed by a
digital camera fitted to the stereomicroscope. A few fruiting
bodies were transferred to a drop of water mounted on a slide
using a fine needle or forceps for rehydration and they were
crushed to show internal structures. Hand sections of fruit
bodies were made by a razor blade and mounted in a water
drop. Morphological characteristics of ascomata/conidiomata,
asci, ascospores/conidia and other tissues associated with
ascomata were photographed using a digital camera fitted with
the compound microscope. Melzer’s reagent, Indian ink and
5 % KOH were used when necessary. All microphotographs
were arranged using Adobe Photoshop CS3 extended (v. 10.0)
and all measurements were made with Tarosoft image
Fungal Diversity
framework (v. 0.9.0.7). Specimens were preserved and deposited at accessible herbaria.
Single spore isolation
Few fruiting bodies were picked from the substrate using a
sterilized needle and placed in few drops of sterilized distilled
water mounted on a sterilized cavity slide as detailed in
Chomnunti et al. (2014). Fruiting bodies were crushed to separate the spores and obtain a spore suspension. The suspension
was inoculated on to fresh Malt Extract Agar (MEA) or Potato
Dextrose Agar (PDA) plates and incubated at room temperature. Germinating ascospores/conidia were aseptically transferred to fresh culture plates. Colonies were photographed and
characters noted. Living cultures are deposited at BCC,
MFLU, GU and ICMP culture collections.
Promoting asexual morphs
Dried and sterilized, grass leaves or pine needles were laid
parallel to the colony margin of pure cultures which were
grown on water agar. Conidiomata produced on both water
agar and sterilized pine needles were observed after few weeks
of incubation at 20 °C. Morphological characteristics were
studied as mentioned above.
Herbarium specimen examination
Types and other relevant authentic specimens were loaned
from different accessible herbaria (G, MFLU, NY, PDD, S,
WU). A small part of the herbarium specimen was cut and
rehydrated using 5 % KOH. Micro-morphological characters
were observed from rehydrated ascomata and photography
was done as previously described. Some specimens were redrawn based on previously illustrated authentic specimens.
DNA extraction, PCR amplification, sequencing
and phylogenetic analyses
Fresh fungal mycelium grown on MEA or PDA for 4 weeks at
20 °C was scraped from the colony margin and were used for
genomic DNA extraction using a modified protocol of Doyle
and Doyle (1987) and Lee and Taylor (1990). PCR amplification and sequencing of LSU region using the primer pair
LROR/LR5 and ITS region using primer pair ITS5/ITS4
was performed (Vilgalys and Hester 1990; White et al.
1990). The PCR products were visualized on 1 % agarose
electrophoresis gel stained with ethidium bromide.
Purification and sequencing of PCR products were carried
out at BGI Ltd. Shenzhen, PR China. The DNA sequences
generated from this study were deposited at GenBank.
The sequences generated in this study were supplemented
with additional sequences obtained from GenBank (Table 1)
based on blast searches and published literature. Multiple sequence alignments were generated with MAFFT v. 7 (http://
mafft.cbrc.jp/alignment/server/index.html); the alignment was
improved with MEGA v. 5.2.2 (Kumar et al. 2012) or BioEdit
v. 7.0.5.2 (Hall 1999). The phylogenetic analyses were performed for Maximum Likelihood in RAxML 7.4.2 Black Box
or RAxMl GUI (Stamatakis 2006; Stamatakis et al. 2008),
Maximum Parsimony in PAUP v. 4.0b10 (Swofford 2003),
or Bayesian inference in MrBayes v. 3.2.1 (Ronquist et al.
2012) as specified in the legend of each phylogenetic tree.
Results and discussion
The phylogeny resulting from the analysis of LSU and ITS
sequence data is shown in Fig. 1. Overall, the topologies obtained from the different phylogenetic analyses were similar.
The separation between Xylariales and Amphisphaeriales is
not well-supported, however morphologically they are distinct
and we treat them as distinct orders.
The order Amphisphaeriales separates into nine clades labeled as A-H with varying support. Clade A has weak support
and is represented by Discosiaceae fam. nov. Species of
Adisciso, Discosia, Discostroma, Sarcostroma and
Seimatosporium grouped in Clade A in congruence with previous findings (Maharachchikumbura et al. 2014; Li et al.
2015a). Species of Discostroma and Seimatosporium have
been linked by molecular data (Tanaka et al. 2011), however
the molecular links have not been proven for the types of
either genera. Although the support is weak, we introduce this
as a new family because of morphological similarities in the
included genera and the fact that this group of genera always
cluster together in analyses. Clade B includes Ciliochorella,
Lepteutypa, Monochaetia, Neopestalotiopsis, Pestalotiopsis,
Pseudopestalotiopsis and Seiridium and are accommodated
in a new family, Pestalotiopsidaceae introduced in this paper.
Both morphological similarities and sequence analysis support their status as a family. Clade C includes species of
Bartalinia, Broomella, Dyrithiopsis, Hyalotiella, Truncatella
and Zetiasplozna and a new family Bartaliniaceae is introduced to accommodate these taxa. Both morphological similarities and sequence analysis support their establishment as a
family. Clades D include the genera Immersidiscosia and
Ellurema, and are phylogenetically closely related to
Discosiaceae, Pestalotiopsidaceae and Bartaliniaceae.
However, the relationships of Immersidiscosia and Ellurema
to any of the above families are poorly resolved and therefore
the genera are placed in Amphisphaeriales genera incertae
sedis (the LSU region of Ellurema indica isolate MIU43478,
accession: U43478, needs resequencing as it has a short sequence, approximately around 450 bp). Clade E comprises of
two species of the asexual genus Phylogicylindria and two
species of the sexual genus Ciferriascosea and these are
Fungal Diversity
Table 1 Isolates sequenced in
this study and their GenBank and
culture accession numbers
Taxon name
Culture accession no
Genbank accession no
LSU
ITS
Amphisphaeria sorbi
MFLUCC 13–0721
KP744475
KR092797
Annulohypoxylon multiforme
Arthrinium hyphopodii
Arthrinium subglobosum
Bartalinia robillardoides
Biscogniauxia nummularia
Broomella vitalbae
Broomella vitalbae
Cainia anthoxanthis
Cainia graminis
Ciferriascosea rectamurum
Ciferriascosea fluctamurum
Diatrype disciformis
Daldinia concentrica
Discosia neofraxinea
Discostroma fuscellum
Hyalotiella rubi
Eutypa flavovirens
Hyalotiella spartii
Kretzschmaria deusta
CBS119016
MFLUCC 15–0003
MFLUCC 11–0397
MFLUCC 12–0070
MUCL 51395
MFLUCC 14–1000
MFLUCC 13–0798
MFLUCC 15–0539
MFLUCC 15–0540
MFLUCC 15–0542
MFLUCC 15–0541
MFLUCC 15–0538
CBS113277
MFLUCC 13–0204
MFLUCC 14–0052
MFLUCC 13–0660
MFLUCC 13–0625
MFLUCC 13–0397
CBS 163.93
KT281893
KR069111
KR069113
KR559738
KT281894
KP757750
KP757749
KR092777
KR092781
KR092776
KR092778
KR092784
KT281895
KR072672
KT005514
KR092775
KR092774
KP757752
KT281896
KC477234
KR069110
KR069112
KT162919
JX658444
KP757754
KR092788
KR092787
KR092793
KR092786
KR092789
KR092795
AY616683
KR072673
KT005515
KR092799
KR092798
KP757756
KT281901
Podosordaria tulasnei
Poronia punctata
Pseudomassaria chondrospora
Pseudomassaria corni
Sarcoxylon compunctum
Seimatosporium cornii
Seimatosporium rhombisporum
CBS 128.80
CBS 656.78
MFLUCC 15–0545
MFLUCC 15–0544
CBS 359.61
MFLUCC 14–0467
MFLUCC 15–0543
KT281897
KT281900
KR092779
–
KT281898
KR559739
KT281902
KT281904
KR092790
KR092791
KT281903
KT162918
Seimatosporium ficeae
Seimatosporium rosae
Seimatosporium vitis
Truncatella spartii
Vialaea mangiferae
Xylaria obovata
Xylaria polymorpha
MFLUCC 15–0519
MFLUCC 14–0621
MFLUCC 14–0051
MFLUCC 15–0537
MFLUCC 12–0808
MFLUCC 13–0115
MUCL 49884
KR092780
KR920686
KT198727
KR920362
KR092782
KF724975
KR049089
KT281899
KR092792
KR092800
KT198726
KR920363
KR092794
KF724974
KR049088
FN689809
accommodated in a new family, Phylogicylindriaceae introduced in this paper (note: the LSU sequences of
Ciferriascosea are rather short; accession: KR092778,
KR092776). Clade F is represented by the family
Clypeosphaeriaceae and is well-separated from
Amphisphaeriaceae sensu stricto. Clade G has good
support and represents Amphisphaeriaceae sensu stricto.
Two Amphisphaeria species are included here. Clade H
is represented by P. carolinensis (IFO 9502), a taxon
that may not belong to Pseudomassaria. The same observation was reported by Jaklitsch and Voglmayr
(2012).
The order Xylariales comprises 14 clades with varying
support labeled I-V. Clade I represents the xylaroid clade of
Xylariaceae which includes the epitype of Xylaria hypoxylon
Fig. 1 Consensus tree resulting from a Maximum Likehood analysis of
the combined LSU and ITS sequence alignment of Xylariomycetidae and
other isolates in Sordariomycetes. Families are indicated in coloured
blocks. RAxML bootstrap support values (MLB above 50 %), Bayesian
posterior probabilities (PP above 90 %) and maximum parsimony
bootstrap support values (MPB above 50 %) are given at the nodes
(MLB/PP /MPB). The scale bar represents the expected number of
changes per site. The tree was rooted to Dothidea sambuci and Botrytis
fuckeliana
Fungal Diversity
77/ 1.0/ 54
Discostroma tostum NBRC 32626
Seimatosporium rosae MFLUCC 14−0621
Seimatosporium cornii MFLUCC 14–0467
66/ 0.91/ 61
Discostroma fuscellum MFLUCC 14–0052
Seimatosporium hypericinum NBRC 32647
78/ 1.0/ 80
Seimatosporium rhombusii MFLUCC 15–0543
100 / 1.0/ 99
Seimatosporium ficeae MFLUCC 15–0519
90/ 1.0/ 82
Discosiaceae
Sarcostroma restionis CBS 118154
77/ 1.0/ 51
Seimatosporium eucalypti CBS 115131
-/ 0.90/ Discosia neofraxinea MFLUCC 13–0204
Adisciso yakushimense MAFF 242774
87/ 1.0/ 92
Adisciso tricellulare NBRC 32705
98/ 1.0/ 98
Discosia pini MAFF 410149
100/ 1.0/ 100
Discosia artocreas NBRC 8975
Lepteutypa cupressi IMI 052255
55/ -/ Ciliochorella castaneae HHUF 28799
52/ 0.92/ Seiridium phylicae CPC 19965
50/ 1.0/ Seiridium cardinale CBS 172.56
Seiridium
papillatum CBS 340.97
53 / Monochaetia kansensis PSHI2004Endo1032
100/ 1.0/ 100
Pestalotiopsaceae
Monochaetia kansensis PSHI2004Endo1030
74/ 0.95/
Pseudopestalotiopsis cocos CBS 272.29
66/ -/ 70
57
Pseudopestalotiopsis theae MFLUCC 12–0055
100/ 1.0/ 100
Neopestalotiopsis protearum CBS 114178
100/ 1.0/ 100
Neopestalotiopsis
rosae CBS 101057
99/ 1.0/ 92
Pestalotiopsis malayana CBS 102220
100/ 1.0/ 100
Pestalotiopsis knightiae CBS 114138
50/ 0.92/ 68
Truncatella spartii MFLUCC 15–0537
82/ -/ 93/ -/ 57 Truncatella spartii MFLUCC 15–0573
67/ -/ - Truncatella sp. HKUCC 7987
88/ -/ Truncatella restionacearum CMW 18755
-/ 1.0/ 84
Truncatella hartigii CBS 118148
Dyrithiopsis lakefuianensis HKUCC 7303
Zetiasplozna acaciae CPC 23421
Bartalinia robillardoides CBS 122705
Bartaliniaceae
-/ -/ 94
Bartalinia robillardoides MFLUCC 12–0070
Hyalotiella rubi MFLUCC 13–0660
Hyalotiella spartii MFLUCC 13–0397
Broomella vitalbae MFLUCC 14–1000
99/ 1.0/
100/ 1.0/ 100
98
Broomella vitalbae MFLUCC 13–0798
100/ 1.0/
100
Truncatella
laurocerasi ICMP 11214
88/ 1.0/ 88
Truncatella angustata ICMP 7062
Ellurema indica MIU 43478
-/ 0.96/ Immersidiscosia eucalypti NBRC 104195
100/ 1.0/ 99
Immersidiscosia eucalypti MAFF 242781
Ciferriascosea wavyseptata MFLUCC 15–0541
80/ 0.98/ Ciferriascosea
uniseptata
MFLUCC 15–0542
78/ 0.98/ 58
54/ 0.98/
Phlogicylindriaceae
Phlogicylindrium eucalyptorum CBS 111689
100/ 1.0/
Phlogicylindrium uniforme CBS 131312
100
Clypeosphaeria uniseptata HKUCC 6349
100/ 1.0/ 100
Clypeosphaeriaceae
Clypeosphaeria
uniseptata AY083830
98/ 1.0/ 95
Amphisphaeria umbrina HKUCC 994
76/ -/ 87
Amphisphaeriaceae
Amphisphaeria sorbi MFLUCC 13–0721
69/ 0.99/ Pseudomassaria carolinensis 9502 (IFO)
87/ 1.0/ 80
a
b
c
d
e
f
g
h
Fungal Diversity
Podosordaria tulasnei CBS 128.80
Xylaria hypoxylon CBS 122620
Xylaria obovata MFLUCC 13–0115
79/ 0.98/ Xylaria polymorpha MUCL 49884
99/ 0.98/ 96
Kretzschmaria deusta CBS 163.93
Poronia
punctata CBS 656.78
52/ 0.92/ 76/ 1.0/ 50
Sarcoxylon compunctum CBS 359.61
Biscogniauxia nummularia MUCL 51395
95/ 1.0/ 91
Camillea obularia ATCC 28093
88/ 0.97/ 91
60/ 0.98/ 66/ -/ -
“Xylarioideae” Xylariaceae
i
67/ 0.96/ -
Graphostroma platystoma AFTOL ID 1249 Graphostromataceae
Hypoxylon
fragiforme MUCL 51264
95/ 1.0/ 77
“Hypoxyloideae” Xylariaceae
Daldinia concentrica CBS 113277
99/ 1.0/ 72
Annulohypoxylon multiforme CBS 119016
100/ 1.0/ 64 Cainia graminis CBS 136.62
-/ 0.98/ 100/ 1.0/ 100
Cainia anthoxanthis MFLUCC 15–0539
92/ 1.0/ 62
Cainia graminis MFLUCC 15–0540
Atrotorquata spartii MFLU 14–0738 Cainiaceae
99/ 0.99/ 79
Seynesia erumpens SMH 1291
Amphibambusa bambusicola MFLUCC 11–0617
-/ -/ 70
Arecophila bambusae HKUCC 4794
Lopadostoma americanum HV-2014h LG8
64/ -/ 84/ 0.99/ 84
Lopadostoma dryophilum LG21
89/ 1.0/ 97
Lopadostoma quercicola HV-2014a strain LG27 Lopadostomaceae
Lopadostoma turgidum LT2
94/ 1.0/ 78
-/ -/ 79
Lopadostoma fagi HV-2014f LF1
Creosphaeria sassafras CM AT-018
-/ -/ 58
Coniocessia nodulisporioides CBS 281.77
100/ 1.0/ 100
Coniocessiaceae
Coniocessia maxima strain Co117
- / 0.98/ Diatrype disciformis MFLUCC 15–0538
-/ 1.0/ 84
Eutypa lata CBS 208.87
Diatrypaceae
Diatrype palmicola MFLUCC 11–0018
97/ -/ 58
Eutypa flavovirens MFLUCC 13–0625
Diatrype whitmanensis ATCC MYA-4417
95/ -/ 63 Apiospora bambusae ICMP 6889
84/ 1.0/ 97
Apiospora setosa ICMP 4207
82/ 1.0/ 89
Arthrinium sp. MFLUCC 15–0002
-/ 0.90/ Apiosporaceae
82/ 1.0/ 95
Arthrinium phaeospermum HKUCC 3395
Arthrinium hyphopodii MFLUCC 15–0003
82/ 0.99/ Arthrinium subglobosum MFLUCC 11–0397
100/ 1.0/
Pseudomassaria chondrospora MFLUCC 15–0545
100
Pseudomassaria chondrospora PC1
Pseudomassariaceae
100/ 1.0/ 100
Pseudomassaria sepincoliformis CBS 129022
Hyponectria buxi UME 31430
Hyponectriaceae
Melogramma campylosporum MBU
Melogrammataceae
Squamotubera sp. HKAS79008
100/ 1.0/ 100
Squamotubera sp. HKAS78918
99/ 0.99/ 100
Iodosphaeria phyllophila FJS8
Iodosphaeriaceae
81/ 0.97/ 79
100/ 1.0/ 97 Repetophragma inflatum NN 42958
Sporidesmium knawiae CPC 15467
Vialaea minutella BRIP 56959
100/ 1.0/ 100
Vialaeaceae
Vialaea mangifia MFLUCC 12–0808
Sordaria fimicola HKUCC 3714
97/ 1.0/ 95
Sordariomycetidae
Apiosordaria verruculosa F-152365
93/ 0.99/ 54
Meliola centellae VIC 31244
100/ 1.0/ 100
69/ 0.98/ Meliolomycetidae
Asteridiella obesa VIC 31239
Cryptodiaporthe aesculi AFTOL-ID 1238
100/ 1.0/ 100
Diaporthomycetidae
Ophiodiaporthe cyatheae YMJ 1364
84/ 0.99/ Hypocrea rufa DAOM JBT1003
Hypocreomycetidae
84/ 1.0/ Reticulascus clavatus CBS 125296
Dothidea sambuci DAOM 231303
Botryotinia fuckeliana AFTOL-ID 59
j
k
l
m
n
o
p
q
r
s
t
u
v
0.2
Fig. 1 (continued)
Fungal Diversity
(CBS 122620), the type species of Xylaria. However the
strains in this subfamily have short LSU sequence data.
Other species groups in the clade I are Podosordaria,
Kretzschmaria, Poronia and Sarcoxylon. Clade J is represented the hypoxyloid clade of Xylariaceae. However, within the
clade J includes the monotypic family Graphostromataceae
represented by the generic type Graphostroma platystoma,
showing affinities with Biscogniauxia Kunze and Camillea
Fr (see notes under Graphostroma). Clade K is the family
Cainiaceae comprising Amphibambusa bambusicola,
Arecophila bambusae, Atrotorquata spartii, Cainia spp. and
Seynesia erumpens. Atrotorquata was previously placed in
Amphispharaceae (Lumbsch and Huhndorf 2007;
Maharachchikumbura et al. 2015) and Kang et al. (1999a, b)
pointed out that it morphologically resembles Cainia. Clade L
represents a newly introduced family Lopadostomaceae.
Generally the obtained data indicate a close relationship between Lopadostoma and Creosphaeria and is in agreement
with Jaklitsch et al. (2014). All Coniocessia LSU sequence
data in GenBank are short and clade M represents the family
Coniocessiaceae. Another significant clade was formed by the
family Diatrypaceae and is represented by clade N as a sister
group to Lopadostomaceae and Coniocessiaceae.
Diatrypaceae comprises Eutypa / Diatrype species. Another
significant clade is the family Apiosporaceae (Clade O) and
Crous and Groenewald (2013) analyzed LSU rDNA sequence
data and confirmed that the Apiosporaceae belongs in
Xylariales, and is a sister group to Amphisphaeriaceae.
Apiosporaceae was a sister group to Pseudomassariaceae in
this study. Clade P corresponds to Pseudomassaria and is
accommodated in a new family, Pseudomassariaceae. Based
on LSU sequence data, Jaklitsch and Voglmayr (2012)
reported anomalies in the taxonomic position of
Hyponectira buxi and they questioned the reliability of
this strain. In the present study it (clade Q) form a sister
clade to family Melogrammataceae. Clade R comprises the
monotypic family Melogrammataceae. The genus
Squamotubera is represented by clade S. Sequences analysis
of Iodosphaeria (clade T) results in a distinct taxon and in the
family Iodosphaeriaceae. Clade U is represented by
Sporidesmium knawiae which is considered as synonymous
to the sister species, Repetophragma inflatum. Clade V comprises of two species of Vialaea in the family Vialaeaceae.
Taxonomy
Outline of Xylariomycetidae
$
=Sequence data in GenBank as at date on submission
Subclass Xylariomycetidae O.E. Erikss & Winka
Order Amphisphaeriales D. Hawksw. & O.E. Erikss.
Amphisphaeriaceae G. Winter
Amphisphaeria Ces & De Not. $
Bartaliniaceae Wijayawardene et al.
Bartalinia Tassi$
Broomella Sacc$
Dyrithiopsis L. Cai et al. $
Hyalotiella Papendorf$
Truncatella Steyaert$
Zetiasplozna Nag Raj $
Clypeosphaeriaceae G. Winter
Apioclypea K.D. Hyde$
Aquasphaeria K.D. Hyde
Brunneiapiospora K.D. Hyde et al. $
Clypeosphaeria Fuckel$
Crassoascus Checa et al.
Palmomyces K.D. Hyde et al. $
Discosiaceae Maharachch. & K.D. Hyde
Adisciso Kaz. Tanaka$
Discosia Lib. $
Discostroma Clem. $
Sarcostroma Cooke$
Seimatosporium Corda$
Pestalotiopsidaceae Maharachch. & K.D. Hyde
Ciliochorella Syd. $
Neopestalotiopsis Maharachch. et al. $
Pestalotiopsis Steyaert$
Pseudopestalotiopsis Maharachch. et al. $
Seiridium Nees$
Phlogicylindriaceae Senan., & K.D. Hyde
Ciferriascosea Senan., et al. $
Phlogicylindrium Crous et al. $
Amphisphaeriales, genera incertae sedis
Amphisphaerella (Sacc.) Kirschst.
Beltraniella Subram.$
Bleptosporium Steyaert
Capsulospora K.D. Hyde$
Ceratosporium Schwein.
Ceriophora Höhn.
Ceriospora Niessl
Chitonospora E. Bommer et al.
Clypeophysalospora H.J. Swart
Distorimula San Martín et al.
Doliomyces Steyaert
Dyrithium M.E. Barr
Ellurema Nag Raj & W.B. Kendr. $
Flagellosphaeria Aptroot
Frondispora K.D. Hyde
Funiliomyces Aptroot$
Griphosphaerioma Höhn.
Hyalotiopsis Punith.
Immersidiscosia Kaz. Tanaka et al. $
Labridium Vestergr.
Lindquistomyces Aramb. et al.
Manokwaria K.D. Hyde
Microdochium Syd. $
Monochaetinula Muthumary et al.
Monographella Petr. $
Morinia Berl. & Bres. $
Fungal Diversity
Mukhakesa Udaiyan & Hosag.
Neobroomella Petr.#
Ommatomyces Kohlm et al.#
Paracainiella Lar.N. Vassiljeva
Pemphidium Mont.
Reticulosphaeria Sivan. & Bahekar
Robillarda Sacc. $
Sporocadus Corda$
Synnemapestaloides T. Handa & Y. Harada
Urosporella G.F. Atk.
Urosporellopsis W.H. Hsieh et al.
Xylochora Arx & E. Müll.
Zetiasplozna Nag Raj$
Xylariales Nannf.
Apiosporaceae K.D. Hyde et al.
Appendicospora K.D. Hyde$
Arthrinium Kunze$
Dictyoarthrinium S. Hughes
Endocalyx Berk. & Broome
Scyphospora L.A. Kantsch.
Spegazzinia Sacc. $
Cainiaceae J.C. Krug
Arecophila K.D. Hyde$
Atrotorquata Kohlm. & Volkm.-Kohlm. $
Cainia Arx & E. Müll. $
Seynesia Sacc. $
Coniocessiaceae Asgari & Zare
Coniocessia Dania García et al. $
Diatrypaceae Nitschke
Anthostoma Nitschke$
Cryptosphaeria Ces & De Not. $
Cryptovalsa Ces & De Not. ex Fuckel$
Diatrype Fr. $
Diatrypella (Ces & De Not.) De Not. $
Diamantinia A.N. Mill. et al. $
Diatrypasimilis J.J. Zhou & Kohlm. $
Echinomyces Rappaz
Eutypa Tul. & C. Tul. $
Eutypella (Nitschke) Sacc. $
Leptoperidia Rappaz
Libertella Desm.
Monosporascus Pollack & Uecker$
Pedumispora K.D. Hyde & E.B.G. Jones$
Peroneutypa Berl. $
Phaeoisaria Höhn. $
Quaternaria Tul. & C. Tul.
Graphostromataceae M.E. Barr et al.
Graphostroma Piroz. $
Hyponectriaceae Petr.
Apiothyrium Petr.
Arecomyces K.D. Hyde
Arwidssonia B. Erikss.
Cesatiella Sacc.
Chamaeascus L. Holm et al.
Charonectria Sacc.
Discosphaerina Höhn. $
Exarmidium P. Karst.
Frondicola K.D. Hyde
Hyponectria Sacc. $
Micronectria Speg.
Papilionovela Aptroot
Pellucida Dulym. et al.
Physalospora Niessl$
Phragmitensis M.K.M. Wong et al.
Rhachidicola K.D. Hyde & J. Fröhl.
Xenothecium Höhn.
Iodosphaeriaceae O. Hilber
Iodosphaeria Samuels$
Lopadostomaceae Daranagama & K.D. Hyde
Creosphaeria Theiss. $
Lopadostoma (Nitschke) Traverso$
Melogrammataceae G. Winter
Melogramma Fr. $
Pseudomassariaceae Senan., & K.D. Hyde
Leiosphaerella Hohn.$
Pseudomassaria Jacz.$
Vialaeaceae P.F. Cannon
Vialaea Sacc. $
Xylariaceae Tul. & C. Tul.
“Hypoxyloideae”
Annulohypoxylon Y.M. Ju et al. $
Anthocanalis Daranagama et al. $
Barrmaelia Rappaz$
Biscogniauxia Kuntze$
Calceomyces Udagawa & S Ueda$
Camillea Fr. $
Chlorostroma A.N. Mill. et al.
Collodiscula I Hino & Katum. $ (cf.Stadler et al. 2013)
Daldinia Ces. & De Not. $
Entonaema Möller$
Hypoxylon Bull. $
Nodulisporium Preuss$
Obolarina Pouzar$
Phylacia Lév. $
Pyrenomyxa Morgan$
Pyriformiascoma Daranagama et al. $
Rhopalostroma D. Hawksw. $
Rostrohypoxylon J Fourn & M Stadler$
Ruwenzoria J. Fourn. et al. $
Thamnomyces Ehrenb. $
Thuemenella Penz. & Sacc. $
Vivantia J.D. Rogers et al.
“Xylarioideae”
Amphirosellinia Y.M. Ju et al. $
Anthostomella Sacc. $
Appendixia B.S. Lu & K.D. Hyde
Fungal Diversity
Areolospora S.C. Jong & E.E. Davis
Arthroxylaria Seifert & W Gams$
Ascotricha Berk. $
Astrocystis Berk. & Broome$
Brunneiperidium Daranagama et al. $
Cannonia J.E. Taylor & K.D. Hyde#
Chaenocarpus Rebent.#
Coniolariella Dania García et al. $
Cyanopulvis J. Fröhl. & K.D. Hyde
Durotheca Læssøe et al.
Emarcea Duong et al. $
Engleromyces Henn.
Entoleuca Syd. $
Euepixylon Füisting$
Fasciatispora K.D. Hyde$
Gigantospora B.S. Lu & K.D. Hyde
Guestia G.J.D. Sm. & K.D. Hyde$
Halorosellinia Whalley et al. $
Helicogermslita Lodha & D. Hawksw.
Hypocopra (Fr) J. Kickx f$
Hypocreodendron Henn.
Induratia Samuels et al. $
Jumillera J.D. Rogers et al.
Kretzschmaria Fr. $
Kretzschmariella Viégas
Leprieuria Laessøe et al.
Leptomassaria Petr.
Lunatiannulus Daranagama et al. $
Muscodor Worapong et al. $
Myconeesia Kirschst.
Nemania Gray$
Nipicola K.D. Hyde
Occultitheca J.D. Rogers & Y.M. Ju$
Ophiorosellinia J.D. Rogers et al. $
Pandanicola K.D. Hyde
Paramphisphaeria F.A. Fernández et al.
Paucithecium Lloyd
Podosordaria Ellis. & Holw. $
Poroleprieuria M.C. González et al.
Poronia Willd. $
Rosellinia De Not. $
Sabalicola K.D. Hyde
Sarcoxylon Cooke
Sclerodermatopsis Torrend
Spirodecospora B.S. Lu et al.
Steganopycnis Syd. & P Syd.
Stilbohypoxylon Henn. $
Streblema Chevall.
Striatodecospora D.Q. Zhou et al. $
Stromatoneurospora S.C. Jong & E.E. Davis$
Theissenia Maubl. $
Virgaria Nees$
Wawelia Namysl.
Xylaria Hill ex Schrank$
Xylocrea Möller
Xylotumulus J.D. Rogers et al. $
Xylariales, genera incertae sedis
Adomia S. Schatz
Ascotrichella Valldos. & Guarro
Biporispora J.D. Rogers et al.
Diamantinia A.N. Mill. et al. $
Fassia Dennis
Lanceispora Nakagiri et al. $
Lasiobertia Sivan. $
Leiosphaerella Höhn.$
Linocarpon Syd. & P. Syd. $
Neolinocarpon K.D. Hyde$
Oxydothis Penz. & Sacc. $
Palmicola K.D. Hyde
Pidoplitchkoviella Kiril. $
Plectosphaera Theiss. $
Polyancora Voglmayr & Yule$
Pulmosphaeria Joanne E. Taylor et al. $
Sporidesmina Subram. & Bhat$
Squamotubera Henn.
Subramaniomyces Varghese & V.G. Rao$
Yuea O.E. Erikss.
Xylariomycetidae genera incertae sedis
Neohypodiscus J.D. Rogers et al.#
Whalleya J.D. Rogers et al. $
The latest outline of Xylariomycetidae was published by
Maharachchikumbura et al. (2015). However, the present
study, Norphanphoun et al. (2015) and Li et al. (2015b) have
revised the group. A new outline is therefore provided below.
This should be regarded as a working outline as there are
many genera that lack sequence data, others where the type
species has not been sequenced, some with incomplete sequence data, and other genera with only one or two taxa
sequenced.
The subclass Xylariomycetidae was introduced by
Eriksson and Winka (1997) and many species in the
subclass have large stromata, containing numerous
ascomata, asci with a J+ apical apparatus and brown
unicellular ascospores (Maharachchikumbura et al. 2015).
However, they may also have single ascomata, often
immersed under a clypeus, and unicellular, apiosporous,
trans-septate or muriform ascospores, ranging from
hyaline to black (Zhang et al. 2006; Stadler et al. 2013).
The subclass is presently rather confusing as many,
especially the astromatic genera, lack sequence data
(Maharachchikumbura et al. 2015). In addition, the connections of the sexual and asexual morphs have not been
established for many of the stromatic genera, in particular
in the Xylariaceae.
Amphisphaeriales D. Hawksw. & O.E. Erikss., Syst.
Ascom. 5 (1): 177 (1986)
Fungal Diversity
Facesoffungi number: FoF 00672
Saprobic, pathogenic or endophytic on plants, appearing as
black spots on the host substrate, made up of fungal hyphae
and host cells. Sexual morph: Pseudostromata or stromata if
present, with immersed to erumpent, solitary, rarely aggregated, scattered, globose to subglobose, coriaceous, brown to
black, ostiolate, ascomata with periphyses. Peridium with
one or few layers, mostly comprising an outer layer of thickwalled, brown cells and inner layer of thick or thin-walled,
hyaline cells. Hamathecium comprising septate, hyaline, paraphyses. Asci 8-spored, unitunicate, cylindrical, clavate or
narrowly globose, short pedicellate, with J+ or J- apical ring.
Ascospores uniseriate or multi-seriate, hyaline, light brown to
dark brown, ellipsoid, apiosporous, fusiform or oval, unicellular to 4-septate smooth or ornamented, sometimes bearing
hyaline, slender, unbranched appendages, sometimes with a
gelatinous sheath. Asexual morph: Coelomycetous.
Conidiomata stromatic, solitary to gregarious, immersed to
erumpent, acervular, pycnidial to irregular, unilocular or irregularly plurilocular, dark brown to black. Peridium comprising
brown cells of textura angularis. Conidiophores indistinct or
if distinct, arising from the inner layer cells of basal stroma,
sometimes reduced to conidiogenous cells, long, cylindrical,
hyaline, septate and sometimes branched. Conidiogenous
cells discrete, holoblastic, annelledic, indeterminate, integrated, cylindrical, ampulliform, or lageniform, smooth, hyaline,
becoming pale brown with age. Conidia cylindrical, clavate,
ellipsoid or fusiform, aseptate to 4-euseptate, hyaline,
subhyaline, pale olivaceous or brown, bearing cellular, filiform or attenuated, sometimes only apical or having both apical and basal appendages.
Notes: The order Amphisphaeriales was introduced by
Eriksson and Hawksworth (1986) to accommodate fungal
taxa in Amphisphaeriaceae, Cainiaceae,
Clypeosphaeriaceae and Hyponectriaceae. Eriksson and
H a w k s w o r t h ( 1 9 8 7 ) h o w e v e r, s y n o n y m i z e d t h e
Amphisphaeriales under Xylariales and this has been followed by subsequent authors (Lumbsch and Huhndorf 2007,
2010). As a result of combined LSU and ITS gene sequence
analysis in this study (Fig. 1), we resurrect the order
Amphisphaeriales with the currently existing families
Amphisphaeriaceae, Clypeosphaeriaceae and four novel families; Bartaliniaceae, Discosiaceae, Pestalotiopsidaceae and
Phlogicylindriaceae. Members in this order show a wide variation in morphology. However the sexual morph of this order
is characterized by immersed to erumpent, mostly solitary,
globose to subglobose, coriaceous ascomata, with or without
stromatic tissues, cylindrical or clavate asci, with J+ or Japical rings and ellipsoid, apiosporous, fusiform or oval, unicellular to trans-septate, variously coloured ascospores, with
smooth or ornamented walls and with presence or absence of
appendages. The asexual morphs have acervular or pycnidial
conidiomata with holoblastic, annellidic, ampulliform, or
lageniform, conidiogenous cells and aseptate to transeuseptate, variously coloured conidia, often bearing
appendages.
Amphisphaeriaceae G. Winter [as ‘Amphisphaerieae’],
Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 259 (1885)
Facesoffungi number: FoF 00673
Saprobic on branches and wood, appearing as slightly
raised, black spots on host surface, often surrounded by black
patches, herein termed the pseudostromata. Sexual morph:
Pseudostromata when present made up of host cells and
brown to black fungal hyphae. Ascomata scattered, solitary
or clustered, immersed, globose to subglobose, coriaceous,
dark brown, ostiolate, papillate. Papilla short, internally covered by hyaline, filamentous periphyses. Peridium two-layered, outer layer of thick-walled, brown cells of textura
angularis, inner layer of small, thin-walled, hyaline cells of
textura angularis. Hamathecium comprising numerous, septate, hyphae-like paraphyses. Asci 8-spored, unitunicate, cylindrical, short pedicellate, with J+ or J- apical ring.
Ascospores uniseriate to overlapping uniseriate, light to dark
brown, oval to fusiform, 1-septate, not or slightly constricted
at the septum, smooth-walled, surrounded by mucilaginous
sheath. Asexual morph: Coelomycetous. Conidiomata globose, dark brown. Conidiophores septate, branched, hyaline.
Conidiogenesis cells elongate conical, septate, annelledic.
Conidia 1-celled elongate-fusiform, hyaline.
Type: Amphisphaeria Ces. & De Not., Comm. Soc. crittog.
Ital. 1 (4): 223 (1863)
Type species: Amphisphaeria umbrina (Fr.) De Not., Sfer.
Ital.: 69 (1863)
Notes: The family Amphisphaeriaceae was introduced by
Winter (1884–1886) to accommodate Amphisphaeria which
is typified by A. umbrina. This family is characterised by
immersed or erumpent perithecial ascomata, some forming
in pseudostromata, with papillate ostioles, unitunicate, cylindrical, asci with J- or J+ apical rings, and light to dark brown,
1-celled ascospores (Kang et al. 1999b). This family was recognized as a heterogeneous sexual morph family accommodating 36 genera (Hawksworth et al. 1995). Kang et al. (1998)
restricted the family to ten genera based on phylogenetic analysis of ITS sequence data, often with pestalotiopsis-like asexual morphs. However, later the family was reduced to 9 genera
(Kirk et al. 2001) excluding some genera to Cainiaceae,
Clypeosphaeriaceae and Hyponectriaceae. In this study we
exclude most genera and restrict the family to
Amphisphaeria sensu stricto.
Type: Amphisphaeria Ces. & De Not., Comm. Soc. crittog.
Ital. 1 (4): 223 (1863)
Saprobic on woody branches, appearing as slightly raised,
black dots on host surface, often surrounded by black patches,
the pseudostromata. Sexual morph: Pseudostromata when
present made up of host cells and brown to black fungal hyphae. Ascomata perithecial, scattered, solitary or clustered,
Fungal Diversity
Fig. 2 Amphisphaeria sorbi (holotype) a Ascomata on substrate. b Cross
section of ascoma. c Peridium. d Paraphyses. e, f Asci in water. g Asci in
Melzer’s reagent. h, i Ascospores. j Sheath around ascospore. k
Germinating ascospore. i Conidiomata on MEA. m–p Conidiophore
and conidiogenous cells with attached conidia. q. Conidia. Scale bars:
a=1 mm, b=100 μm, c=50 μm, e–g=20 μm, d, h–k=10 μm, l–q=
10 μm
immersed, globose to subglobose, coriaceous, dark brown,
ostiolate. papillate. Papilla short, narrow, conical, periphysate.
Periphyses hyaline, filamentous. Peridium two-layered, outer
layer comprising dark brown cells arranged in a textura
angularis and inner layer comprising thin-walled, hyaline
cells. Hamathecium comprising numerous, filamentous,
Fungal Diversity
Fungal Diversity
Bartalinia robillardoides (MFLU 13–0084). a Conidiomata on
leaf of Eucalyptus sp. b c Cross section of conidioma on leaf. d, f
Developing conidia attached to conidiogenous cells. e, f Cross section
of conidioma on PDA. g–l. Conidia. j. Culture on PDA. k. Germinating
conidium. Scale bars: b–e=150 μm, d–f=20 μm, g–i, l=25 μm
Fig. 3
septate, slightly tapering paraphyses. Asci 8-spored,
unitunicate, cylindrical, indistinctly pedicellate, apex rounded,
with J+ or J- apical ring. Ascospores uniseriate to overlapping
uniseriate, light brown to dark brown, oval to fusiform, 1septate, not or slightly constricted at the septum, smoothwalled or ornamented, sometimes with a mucilaginous sheath.
Asexual morph: Coelomycetous. Conidiomata solitary or aggregated, globose, dark brown. Peridium comprising of thickwalled, septate, brown mycelium. Conidiophores septate,
branched, thick-walled, hyaline. Conidiogenous cells elongate
conical, thin-walled, septate, hyaline, annelledic. Conidia hyaline, elongate-fusiform, 1-celled, smooth-walled.
Notes: Amphisphaeria was introduced by Cesati and De
Notaris (1863) without designating a generic type (Wang
et al. 2004). Petrak (1923) proposed A. umbrina as the lectotype of the genus which was introduced earlier as Sphaeria
umbrina. A description of A. umbrina was provided by Hyde
et al. (1996). However this was not based on the type specimen. Hence, Hyde et al. (1996) epitypified a collection listed
by Cesati and De Notaris (1863) as S. umbrina and also used a
specimen used by Petrak (1923) to describe A. umbrina
(Wang et al. 2004). The epitype of A. umbrina is illustrated
in Wang et al. (2004). Amphisphaeria umbrina has a distinct
clypeus comprising outer, brown and inner, hyaline cells of
textura angularis, cylindrical asci, with a J , discoid, subapical
ring and ellipsoidal, light brown, one-median septate ascospores. Wang et al. (2004) accepted 12 species in the genus.
Four species of Amphisphaeria have J-, apical rings (Wang
et al. 2004; Liu et al. 2015). In this study, we illustrate the
sexual and asexual morph of Amphisphaeria sorbi, the latter
being introduced in Lui et al. (2015).
Amphisphaeria sorbi Senan. & K.D. Hyde, in Liu et al.,
Fungal Divers 71: 10 (2015), Fig. 2
Notes: This species was introduced in Liu et al. (2015) and
is similar to A. umbrina. As this species formed an asexual
morph in culture we illustrate both sexual and asexual morphs
here (Fig. 2).
Material examined: ITALY, Trento [TN], Dimaro,
Folgarida, on branch of Sorbus aucuparia L. (Rosaceae), 2
August 2013, E. Camporesi, IT 1400 (MFLU 14–0797,
holotype).
Bartaliniaceae Wijayawardene, Maharachch. & K.D.
Hyde, fam. nov.
Index Fungorum number: IF511183; Facesoffungi
number: FoF 00667
Saprobic or rarely pathogenic causing leaf spots, on various hosts. Sexual morph: Pseudostromata immersed,
solitary, scattered. Ascomata immersed to semi-immersed,
solitary to gregarious, globose to subglobose, black. Ostiole
papillate, central, circular, periphysate. Peridium composed of
brown to dark brown cells of textura angularis or textura
prismatica, with thin inner layer composed of thin-walled,
hyaline cells of textura angularis. Hamathecium comprising
filiform, septate, hyaline, smooth-walled paraphyses. Asci 8spored, unitunicate, cylindrical, short pedicellate, apically
rounded a with J- or J+ subapical ring. Ascospores overlapping uniseriate, pale brown to brown, ellipsoid to fusiform, 3–
4-septate, smooth-walled. Asexual morph: Coelomycetous.
Conidiomata acervular or pycnidial to irregular, solitary to
gregarious, superficial to sub-immersed, unilocular, globose
to subglobose, dark brown to black. Ostiole apapillate.
Conidiomata wall comprising two strata, inner wall composed
of thick-walled, dark brown cells of textura angularis, inner
wall thin, composed of hyaline to sub-hyaline cells of textura
an gula ris. Co nidio pho re s pr esen t or redu ced to
conidiogenous cells; when present cylindrical, hyaline,
sparsely septate, smooth-walled. Conidiogenous cells holoblastic, ampulliform, integrated or discrete, determinate, hyaline, smooth-walled. Conidia fusiform, straight to slightly
curved, subhyaline to brown, bearing only apical appendages
or having both apical and basal appendages.
Type: Bartalinia Tassi, Bulletin Labor. Orto Bot. de R.
Univ. Siena 3: 4 (1900)
Notes: In the molecular analyses (Fig. 1), Bartalinia sensu
stricto groups with Broomella, Hyalotiella and Truncatella
and forms a distinct clade in Amphisphaeriales. Further, the
Bartalinia clade comprises a newly introduced genus viz.
Zetiasplozna. This clade is distinct from the other families in
Amphisphariales and thus we introduce a new family
Bartaliniaceae.
The sexual morph genera in Bartaliniaceae are
characterised by immersed to semi-immersed ascomata with
papillate ostioles, unitunicate, cylindrical asci, with J- or J+
apical rings, and light to dark brown, 3–4-septate ascospores.
The asexual typified genera have superficial to subimmersed
acervular or pycnidial to irregular fruiting bodies, apapillate
ostioles, conidiophores reduced to conidiogenous cells, holoblastic conidiogenesis and fusiform, straight to slightly
curved, subhyaline to brown conidia with apical or apical
and basal appendages.
Bartalinia Tassi, Bulletin Labor. Orto Bot. de R. Univ.
Siena 3: 4 (1900)
Facesoffungi number: FoF 00659
Endophytic or saprobic on substrates of a range of plant
hosts. Sexual morph: Undetermined. Asexual morph:
Conidiomata pycnidial or variable, solitary to gregarious, subepidermal, erumpent at maturity, globose, unilocular, brown to
black. Conidiomata wall two layered, outer layer comprising
dark brown to black cells of textura angularis, inner wall
comprising thin, hyaline cells of textura angularis.
Fungal Diversity
C o n i d i o p h o re s r e d u c e d t o c o n i d i o g e n o u s c e l l s .
Conidiogenous cells ampulliform, holoblastic, discrete, determinate, hyaline, generated from inner layer of the pycnidium wall. Conidia hyaline to pale brown,
subcylindrical, trans-septate, with appendages, basal cell
with truncate base, obconic, hyaline, with single unbranched appendage; 3 median cells subcylindrical, hyaline
to pale brown, apical cell conical, almost hyaline, with threebranched appendage.
Type species: Bartalinia robillardoides Tassi, Bulletin
Labor. Orto Bot. de R. Univ. Siena 3: 4 (1900)
Notes: Tassi (1900) established Bartalinia, with
B. robillardoides Tassi as the type species and currently, the
genus comprises of 19 species epithets (Index Fungorum
2015). Morgan-Jones et al. (1972) accepted only nine species
under Bartalinia and Sutton (1980) also accepted the same
number. Sutton (1980) mentioned that the genus needs taxonomic revision. Von Arx (1981) reduced Bartalinia under
Seimatosporium Corda, but this was rejected by Nag Raj
(1993). Further, Nag Raj (1993) accepted only six species
(including the type species), viz. Bartalinia bischofiae Nag
Raj, B. lateripes (Ellis & Everh.) Nag Raj, B. laurina
(Mont.) Nag Raj, B. pistacina (J.L. Maas) Nag Raj and
B. tamarindi Nag Raj. Five new species have been described
since the revision of Nag Raj (1993) viz. Bartalinia ananatis
Li Zeng et al. (Chi et al. 2002), B. dracaenae P.G. Xi et al. (Xi
et al. 2000), B. goniolimonis Andrian. & Minter (Andrianova
and Minter 2007), B. mellea F. Anderson & Bianchin.
(Anderson and Bianchinotti 1996) and B. pondoensis
Marincowitz et al. (Marincowitz et al. 2010).
Bartalinia robillardoides Tassi, Bulletin Labor. Orto Bot.
de R. Univ. Siena 3: 4 (1900)
≡ Seimatosporium robillardoides (Tassi) Arx, Gen. Fungi
Sporul. Cult., Edn 3 (Vaduz): 224 (1981)
Facesoffungi number; F0F 00660; Fig. 3
Saprobic on leaf, stems and branches of a range of host
plants. Sexual morph: Undetermined. Asexual morph:
Conidiomata 170–240μm diam., 150–200μm high, pycnidial
or variable, solitary to gregarious, of subepidermal in origin,
then erumpent, globose, unilocular, brown to black, with
apapillate ostiole. Pycnidial wall of two layers, outer layer
comprising dark brown to black cells of textura angularis,
inner wall thin, hyaline. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells 2–7 × 2–3 μm,
ampulliform, holoblastic, integrated, determinate, hyaline,
smooth, generated from innermost layer of the pycnidium
wall. Conidia 20–30×3–4μm (x =22.52×3.21μm, n=20),
hyaline to pale brown, subcylindrical, 4-septate, slightly constricted at the septa, smooth-walled, with appendages, basal
cell hyaline, with a truncate base, obconic, 2–4μm long, baring an unbranched single appendage; 3 median cells
subcylindrical, hyaline to pale brown, 15–22 μm long
(altogether); apical cell conical, almost hyaline, 2–4μm long,
with unbranched appendage, attenuate, flexuous, 15–20μm,
tri-radiate.
Culture characteristics: On PDA greyish brown, not
zonate, fast growing, attaining 3 cm in 3 days at 25 °C, comprising circular, flat, thin mycelium.
Material examined: THAILAND, Chiang Rai, Mae Fah
Luang University grounds, on leaves of Eucalyptus sp., 30
June 2012, Nalin N. Wijayawardene, NNW 120630–1
(MFLU 13–0084), living culture, MFLUCC 12–0070,
GUCC T70
Notes: Our collection on leaves of Eucalyptus spp.
morphologically resembles Bartalinia robillardoides
(Nag Raj 1993; Crous et al. 2014a). Further, molecular
data also showed that the new strain resides in
Bartalinia sensu stricto with B. robillardoides (Fig. 3).
Crous et al. (2014a) designated the lectotype and epitype of
B. robillardoides as MBT178268 and CBS H-21728,
respectively.
Broomella Sacc., Syll. fung. (Abellini) 2: 557 (1883)
= Keissleria Höhn., Annls mycol. 16 (1/2): 93 (1918)
Facesoffungi number: FoF 00626
Saprobic on various hosts. Sexual morph: Ascomata solitary to gregarious, uniloculate, glabrous, globose to
subglobose, papillate. Ostiole centrally located, composed of
longitudinally aligned cells, and internally lined with hyaline
periphyses. Peridium composed of light yellow, thick-walled
cells of textura prismatica in the upper part and surrounding
the ostiole, and thin-walled, hyaline to pale brown cells in
other parts. Hamathecium comprising numerous, cylindrical,
hypha-like, septate, paraphyses tapering towards the ends.
Asci 8-spored, unitunicate, cylindrical to cylindrical-clavate,
pedicellate, apically rounded, with a J- apical ring. Ascospores
biseriate or overlapping bi- or tri-seriate, pale greyish brown,
or brown, with ends cells yellowish to pale grayish brown,
fusiform, glabrous, straight or inequilaterally curved, 3septate, constricted at the septa, thick-walled, with
doliform median cells, end cells conical, each bearing
an unbranched, terminal tubular appendage. Asexual morph:
Coelomycetous. Conidiomata stromatic, pycnidioid, scattered
to gregarious, immersed to semi-immersed, rounded, oval or
elongated, black, unilocular, papillate, glabrous. Pycnidial wall
comprising thick-walled cells of textura globulosa to textura
angularis, outwardly pale brown to brown, inwardly merging
with relatively thin-walled and colourless cells. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells integrated,
cylindrical, phialidic, percurrently proliferating 1–2-times, hyaline, smooth. Conidia pale brown or brown, fusiform to
aciculate, with acute ends, straight or slightly curved, 3-septate,
constricted at septa, verruculose, thick-walled, bearing a single
tubular appendage at each end, or 2–5-appendage at apex and a
single appendage at the base.
Type species: Broomella vitalbae (Berk. & Broome) Sacc.,
Syll. fung. (Abellini) 2: 558 (1883)
Fungal Diversity
≡ Hypocrea vitalbae Berk. & Broome, Ann. Mag. nat.
Hist., Ser. 33: 363 (1859)
Facesoffungi number: FoF 00589, Fig. 4
Notes: Broomella was introduced by Saccardo (1883)
based on B. vitalbae (Berk. & Broome) Sacc. The genus is
characterized by unitunicate, cylindrical-elongate asci, with a
Fig. 4 Broomella vitalbae (epitype) a, b Black ascomata on the host. c
Section of peridium. d, e Vertical section of ascomata. f Ostiole. g
Paraphyses. h–k Asci with ascospores. l–o Ascospores. p–q Black
conidiomata on the host. r Vertical section of conidiomata. s
Conidiophores, conidiogenous cells and developing conidia. t–v
Conidia. Scale bars: a=500 μm, b=200 μm, c=20 μm, d–e=100 μm,
f=20 μm, g–o =10 μm, p=200 μm, q=500 μm, r=100 μm, s–v=10 μm
Fungal Diversity
J-, discoid apical ring, and 3-septate, ellipsoid-fusiform ascospores, with brown median cells, lighter terminal cells, and
single, centric appendages arising from the ends (Shoemaker
and Müller 1963; Li et al. 2015a). Broomella has been shown
to be linked to a pestalotiod-like asexual morph (Shoemaker
et al. 1989; Yuan and Zhao 1992; Kang et al. 1999a).
Shoemaker and Müller (1963) introduced B. acuta
Shoemaker & E. Müll. and B. excelsa Shoemaker & E.
Müll. in the genus. There are 20 species epithets listed under
Broomella in Index Fungorum (2015), however, none of these
had been studied using sequence data. The Broomella species
and their truncatella-like asexual morphs differ in various
ways from the type species of Broomella and its asexual
morph and are probably not congeneric. Through a study of
LSU sequence data and together with existing morphological
data, Li et al. (2015a) showed that Broomella appears to be a
distinct, natural group in Amphisphaeriaceae and the asexual
morph have conidia with a single apical and basal
appendages.
Materials examined: ITALY. Province of Forlì-Cesena
[FC], Modigliana, Montebello, on dead stem of Clematis
vitalba, 23 February 2013, Erio Camporesi, IT-1079 (MFLU
15–0065, epitype); ex-epitype living culture, MFLUCC 13–
0 79 8 , I C MP ; I TA LY. P r o vi nc e o f A r e z zo [ A R ] ,
Montemezzano, on dead stem of Clematis vitalba, 25
August 2013, Erio Camporesi, IT-1430 (MFLU 15–0054);
living culture, MFLUCC 14–1000.
Hyalotiella Papendorf, Trans. Br. mycol. Soc. 50 (1): 69
(1967)
Index Fungorum number: IF 8588 Facesoffunginumber:
FoF00627
Saprobic on various hosts. Sexual morph: Ascomata perithecial, immersed, becoming erumpent, solitary, gregarious or
confluent, globose to subglobose, blackened, glabrous, papillate. Ostiole lined with filiform, hyaline, periphyses. Peridium
composed of outer, dark brown, thick-walled cells of textura
angularis and thin-walled, hyaline cells in the inner layers.
Hamathecium comprising filiform, hyaline, paraphyses, evanescent at maturity. Asci 8-spored, unitunicate, cylindrical to
clavate, short pedicellate, apex rounded, with a J+ subapical
ring. Ascospores overlapping uniseriate, fusiform to ellipsoidal with obtuse apex, and with narrow base, initially hyaline,
and becoming brown at maturity, mostly 1-septate, occasionally 2–3-septate, constricted at septa, straight or slightly
curved, thick-walled, smooth. Asexual morph:
Coelomycetous. Conidiomata stromatic, pycnidial, vaseshaped, scattered to gregarious, immersed or semi-immersed,
globose to subglobose, unilocular, glabrous, brown to dark
brown, ostiolate. Ostiole lageniform, single, centrally located,
with a well-developed neck, thick-walled. Peridium thickwalled cells of textura angularis, brown to dark brown.
Conidiophores arising all around the cavity of the conidioma,
short, often reduced to conidiogenous cells. Conidiogenous
cells integrated, cylindrical, long, phialidic, hyaline, smooth.
Conidia cylindrical to fusiform, with an obtuse apex and a
narrow truncate base, straight or slightly curved, 3-septate,
pale brown to brown except for the hyaline apical cell, median
cells longer than ends cells, bearing 2–5-branched, filiform,
flexuous, hyaline appendages arising from apical cell.
Type species: Hyalotiella transvalensis Papendorf, Trans.
Br. mycol. Soc. 50 (1): 69 (1967).
Notes: Hyalotiella was introduced by Papendorf and DuToit (1967) with H. transvalensis Papendorf as type species.
The genus is characterized by having vase-shaped pycnidia,
with a long neck, and 3-septate, cylindrical, pale brown to
brown conidia, with median cells that are longer than the ends
cells, and 2–5-branched, filiform, apical appendages (Nag Raj
1993). According to the Index Fungorum (2015) there are four
epithets in Hyalotiella. However, H. subramanianii was transferred to Hyalotiopsis by Nag Raj (1993) based on conidial
characters and Hyalotiella. orientalis was synonymised under
Hyalotiella americana (Speg.) Nag Raj (Nag Raj 1993). Li
et al. (2015a) introduced an additional taxon in Hyalotiella,
namely, H. spartii W.J. Li et al. based on morphological characters, while they also provided a sequence for this species.
Thus, to date, four species are accepted in Hyalotiella, viz.
H. americana, H. spartii W.J. Li, Camporesi & K.D. Hyde,
H. transvalensis and the sexual species H. rubi described in
this paper.
The Dyrithiopsis-like sexual morph of Hyalotiella has not
been previously reported (Sutton 1980; Nag Raj 1993).
Hyalotiella rubi shows somewhat similar morphological characters (i.e., form of ascomata and ascospores) with the type
species of Dyrithiopsis, viz. Dyrithiopsis lakefuxianensis L.
Cai et al., but it is phylogenetically distinct from
D. lakefuxianensis (Fig. 1) the latter has a Monochaetiopsis
conidial state (Jeewon et al. 2003a, b).
Hyalotiella spartii W.J. Li, Camporesi & K.D. Hyde
Cryptog., Mycol. 36 (1): 93–108 (2015).
Index Fungorum number: IF551047 Facesoffunginumber:
FoF00590; Fig. 5
Saprobic on dead stems of Spartium junceum, forming
conspicuous rounded, black conidiomata. Sexual morph:
Undete rmin ed. Asex ual mor ph: co elom yce to us.
Conidiomata 250–300 μm high, 200–250 μm diam.,
stromatic, pycnidial, vase-shaped, scattered to gregarious, epidermal to subepidermal in origin, globose to subglobose,
semi-immersed, unilocular with the locule occasionally convoluted or irregularly divided, glabrous, brown to dark brown,
ostiolate. Ostiole lageniform, single, centrally located, with a
well-developed neck, thick-walled. Peridium 30–50μm wide,
composed of 7–8-cell layers, with thick-walled cells of textura
angularis, brown to dark brown. Conidiophores arising all
around the cavity of the conidioma, short, often reduced to
conidiogenous cells. Conidiogenous cells 10–15μm wide, integrated, cylindrical, long, phialidic, hyaline, smooth. Conidia
Fungal Diversity
Fig. 5 Hyalotiella spartii (holotype) a Herbarium specimen. b, c Black
coniodiomata on the host. d Vertical section of conidioma. e Ostiole. f, h–j
Conidiogenous cells and developing conidia. g Section of pycnidial wall.
k–n Conidia. Scale bars: c=200 μm, d=100 μm, e=50 μm, f, h–j=10 μm,
g=10 μm, k–n=10 μm
Fungal Diversity
25–28×3–4μm (x =26×3.5μm; n=20), fusiform, 3-septate,
bearing apical appendages; basal cell obconic with an obtuse
base, almost colourless, with 2 cylindrical to subcylindrical,
thick-walled, yellowish to brown, median cells constricted at
septa, and with conic, thin-walled, hyaline apical cell, with
apical appendages 14–20μm long, tubular, acellular, often irregularly or dischotomously branched at the base, filiform,
flexuous, hyaline.
Materials examined: ITALY. Province of Forlì-Cesena
[FC], Santa Sofia, Collina di Pondo, on dead stem of
Spartium junceum, 16 October 2012, Erio Camporesi, IT
812 (MFLU 15–0055 holotype); ex-type living culture,
MFLUCC 13–0397=ICMP 20788; ibid., IT-812B (MFLU
15–0066); living culture MFLUCC 15–0024=ICMP.
Hyalotiella rubi Senan., Camporesi & K. D. Hyde, sp. nov.
Index Fungorum Number: IF551189; Facesoffungi
number: FoF 00680; Fig. 6
Etymology: Name referring to the host genus on which the
species occurs.
Holotype: MFLU 15–0740
Saprobic on stems of Rubus sp., appearing as shiny, black,
raised areas on substrate. Sexual morph: Pseudostromata
made up of host cells and black fungal hyphae, forming a
wide, darkened layer around the ostiole, and a thin, darkened,
hyphal layer beneath the cuticle, spreading on the host surface
to form a darkened region above the ascomata. 140–195μm
high, 200–245μm diam., (x =165×220μm, n=20), perithecial, immersed, becoming erumpent, scattered, solitary or aggregated, globose to subglobose, coriaceous, with very short
papillate ostiole, ostiolar canal lined with filiform, hyaline,
periphyses. Peridium 10–20μm wide (x =16μm, n=20), comprising two strata, outer stratum comprising 7–10 layers of
compressed, thick-walled cells of textura angularis and inner
stratum comprising 1–2 layers of hyaline, thin-walled, large,
cells of textura angularis. Hamathecium comprising 2–5μm
wide (x =3μm, n=20), filiform, septate, hyaline paraphyses,
evanescent at maturity. Asci 80–100×8–10.5μm (x =90×
9.5 μm, n = 20), 8-spored, unitunicate, cylindrical, short
Fig. 6 Hyalotiella rubi (holotype) a Appearance of ascomata on host substrate. b Cross section of ascomata. c Peridium. d Paraphyses. e–g Asci. h Asci
in Melzer’s reagent showing J+, apical ring. i–n Ascospores. o Germinating ascospores. Scale bars: a=1 mm, b=50 μm, c–o=10 μm
Fungal Diversity
pedicellate, apically rounded, with a J+ subapical ring.
Ascospores 12–15μm×5.5–7.5μm (x =13.5×6.5μm, n=20),
overlapping uniseriate, initially hyaline and becoming brown
at maturity, ellipsoidal to fusiform with blunt ends, mostly 1septate, rarely 2–3-septate, slightly constricted at the septa,
lowest cell longer than upper cell, smooth-walled. Asexual
morph: Undetermined.
Culture characters: growing on MEA reaching 2 cm within 7 days when incubated at 18 °C. Flat, circular, white colony
with serrate margin and white woolly centre.
Materials examined: ITALY, Province of Forlì-Cesena
[FC], Premilcuore, Fiumicello, on dead branch of Rubus sp.,
24 April 2013, Nello Camporesi, IT 1197 (MFLU 15–0740,
holotype); Ibid., (MFLU 15–0740bis, isotype); ibid., (MFLU
15–0740tris, paratype); ex-type living cultures, MFLUCC
13–0660=ICMP.
Notes: Hyalotiella rubi is introduced as a new species in the
family Bartaliniaceae based on the phylogenetic analysis of
combined LSU and ITS sequence data. This is the first record
of a sexual morph of Hyalotiella. However Hyalotiella rubi
shows somewhat similar morphological characters to species
of Dyrithiopsis. Phylogenetically they are not closely related.
In the combined analyses of LSU and ITS sequence data of
amphisphaeriaceous and xylariaceous taxa (Fig. 1), H. rubi
shows a close affinity with H. spartii.
Truncatella Steyaert Bull. Jard. bot. État Brux. 19: 293
(1949)
Facesoffungi number: FoF 00705.
Saprobic or mostly pathogenic on plant leaves and twigs.
Sexual morph: Ascomata solitary or gregarious, scattered,
immersed under minute clypeus, uniloculate, globose to
subglobose, coriaceous, papillate or apapillate, brown.
Ostiole central, wide, rounded. Papilla composed of cells of
textura porrecta and internally lined with hyaline periphyses.
Peridium comprising light yellow or brown, thick-walled cells
of textura prismatica in the upper part, and thick-walled, hyaline to pale brown cells of textura angularis in other parts.
Hamathecium comprising numerous, hypha-like, septate, paraphyses, slightly constricted at the septa, tapering towards the
ends. Asci 8-spored, unitunicate, cylindrical to cylindrical-clavate, pedicellate, apically rounded, with a J- apical ring.
Ascospores biseriate or overlapping tri-seriate, hyaline when
young, sometimes pale greyish-brown with doliform median
cells and yellowish to pale greyish-brown end cells or brown
at maturity, fusiform, glabrous, thick-walled, straight or
inequilaterally curved, with pointed ends, 1–3-septate with
constrictions at the septa and bearing unbranched, terminal
appendages or ornamented wall. Asexual morph:
Coelomycetous. Conidiomata acervular, immersed to semiimmersed, solitary to aggregated. Peridium comprising
brown, thin-walled cells of textura angularis. Conidiophores
hyaline, branched, septate, cylindrical. Conidiogenous cells
holoblastic, indeterminate, integrated, cylindrical, hyaline,
smooth, with several percurrent proliferations. Conidia with
brown median cells and hyaline apical and basal cells, fusiform, straight, 3-euseptate, constricted at the septa, sometimes
with a simple or rarely branched, cellular appendage, conical
at apical cell, hyaline, with 1 or more apical, simple or
branched tubular appendages, smooth-walled, thick-walled,.
Type species: Truncatella truncata (Lév.) Steyaert, Bull.
Jard. bot. État Brux. 19: 295 (1949)
Notes: Truncatella introduced by Steyaert (1949) and is
typified by T. truncata. The genus is typically associated with
plants as an endophyte or pathogen (Shoemaker et al. 1989).
According to molecular studies, Truncatella is paraphyletic
with Bartalinia sharing a common ancestor (Jeewon et al.
2002; Li et al. 2015a). However, Truncatella may possibly
be two genera. Truncatella vitalbae (Shoemaker & E. Müll.)
Shoemaker et al., T. excelsa (Petr.) Shoemaker et al., and
T. pestalozzioides (Dearn. & Fairm.) Shoemaker et al. were
reported as the sexual morph of Broomella (Shoemaker et al.
1989). Li et al. (2015a) however, showed through molecular
data that another coelomycetous species is the asexual morph
of B. vitalbae. Hence the sexual morph of Truncatella still
remains unresolved. Here we illustrate the sexual morph of a
new Truncatella species as Truncatella spartii.
Truncatella spartii Senan., Camporesi & K. D. Hyde, sp.
nov. Fig. 6
Index Fungorum number: IF551196; Facesoffungi
number: FoF 00685; Figs. 7 and 8
Etymology: Name referring to the host genus on which the
species occurs.
Holotype: MFLU 15–0721
Saprobic on stems of Spartium junceum L., appearing as
black, circular, flat dots on substrate. Sexual morph:
Ascomata 115–140μm high, 175–190μm diam., (x =135×
180μm, n=20), scattered, solitary, immersed, globose to
subglobose, brown, ostiolate, apapillate. Ostiole wide, circular, even with host epidermis. Peridium 20–24μm wide near
to ostiole (x =22.5μm, n=20), 6.5–13μm wide at the base
(x = 9.5 μm, n =20), outwardly comprising 5–7 layers of
brown cells of textura angularis, inwardly comprising 4–6
layers of thick-walled, hyaline cells of textura angularis.
Hamathecium comprising 2.5–4μm wide (x =3μm, n=20),
branched, septate, paraphyses, slightly constricted at the septa.
Asci 62–70 ×6–6.5 μm (x = 65× 6.1 μm, n= 20), 8-spored,
unitunicate, cylindrical, short pedicellate, apex slightly pointed, with an indistinct, J-, apical apparatus. Ascospores 8–
11.5×3.5–4μm (x =9.5×3.6μm, n=20), uniseriate to overlapping uniseriate, hyaline when young, brown at maturity, fusiform, with one thick septum, constricted at septum, pointed at
ends. Asexual morph: Coelomycetous, Conidiomata 125–
155μm high, 145–180μm diam., (x =140×165μm, n=20),
pycnidial, immersed, g lobose, brown, o stiolate.
Conidiophores 3–5μm high, 3.5–4.5 μm diam. (x =3.5×
4 μm, n = 20), hyaline, simple or branched, cylindrical,
Fungal Diversity
Fig. 7 Truncatella spartii (holotype). a Ascomata on host. b Cross section of ascoma. c Peridium. d Paraphyses. e–i Asci. j–o Ascospores. Scale bars:
a=1 mm, b=50 μm, c, d, j–o=10 μm, e–i=20 μm
aseptate. Conidiogenous cells 8–17μm long, 8–12μm wide,
(x =11.5×10.7μm, n=20), hyaline, cylindrical, annellidic.
Conidia 8–12×4–5.5μm (x =10×5μm, n=20), clavate, 3septate, constricted at the septa, central cells large, upper central cell slightly larger than lower central cell, light brown,
thick-walled, basal cell conical, hyaline, narrowing towards
the base, apical cell small, hyaline, conical, bearing appendages. Appendages 11–16μm long (x =14.5μm, n=20), 1–3,
thin, slender, hyaline, of equal length to that of the conidia.
Culture characters: Growing on MEA reaching 1 cm within 7 days at 18 °C, fast growing, flat, colony circular, with
serrate margin, sparse, white, aerial mycelia in the centre, with
unbranched, hyaline, mycelia at the margin.
Materials examined: ITALY, Province of Forlì-Cesena
[FC], Galeata, Passo delle Forche, on branch of Spartium
junceum L. (Fabaceae), 9 November 2013, Erio Camporesi,
IT 1511 (MFLU 15–0721, holotype); ibid., (MFLU 15–
0721bis, paratype); living cultures, MFLUCC 15–0537
(from conidiomata), ICMP.
Notes: In this study we introduced a new species on
Spartium from Italy as Truncatella spartii. Megablast search
results of T. spartii in GenBank give Truncatella helichrysi
(Severini) B. Sutton and T. hartigii (Tubeuf) Steyaert. as
close matches, however, it differs from them in having
conical, hyaline apical cells, bearing 2–3 long, unbranched, hyaline conidial appendages. We observed
Fungal Diversity
Fig. 8 Truncatella spartii (holotype). a Herbarium specimens. b, c Conidiomata on host. d Cross section of conidioma e–f Conidia attached to
conidiogenous cells. g–l Conidia. Scale bars: b=1 mm, c=200 μm, d=50 μm, e–i=20 μm
ascomata associated with conidiomata on the host and
DNA extraction was made directly from the ascomata
by picking up the inner contents using fine forceps and
checking under a compound microscope. Sequence data
from both the sexual and asexual morphs showed them
to be Truncatella.
Clypeosphaeriaceae G. Winter [as ‘Clypeosphaerieae’],
Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 554 (1886)
Facesoffungi number: FoF 00678
Hemibiotrophic or saprobic on woody or herbaceous
plants. Sexual morph: Pseudoclypeus comprising both host
and fungal tissues, black. Ascomata immersed to erumpent,
rarely superficial, solitary or aggregated, globose to
subglobose, coriaceous, brown to black, ostiolate, papillate.
Papilla short, narrow, internally lined with hyaline, filamentous periphyses. Peridium comprising dark brown to light
brown, thick-walled cells of textura angularis, inwardly hyaline. Hamathecium comprising numerous, hypha-like, septate,
flexuose, paraphyses, embedded in a gelatinous matrix. Asci
8-spored, unitunicate, cylindrical to broadly cylindrical, pedicellate, with a wedge-shaped, J-, or J+ apical ring. Ascospores
uniseriate to biseriate, hyaline to brown, ellipsoidal to fusiform, sometimes oval, straight or curved, unicellular or
septate, wall smooth or ornamented or striate, sometimes with
sheaths, appendages, rarely with germ slits or germ pores.
Asexual morph: Undetermined.
Notes: The family Clypeosphaeriaceae was introduced by
Winter (1884–1886) to accommodate Anthostomella,
Clypeosphaeria, Hypospila, Linospora and Trabutia. Barr
(1989) however, revisited the family and excluded all genera
retaining Clypeosphaeria as the type of this monotypic family.
Hawksworth et al. (1995) treated this family with
Apiorhynchostoma, Ceratostomella, Clypeosphaeria,
Crassoascus, Duradens, Frondicola, Jobellisia, Melomastia
and Pseudovalsaria (Kang et al. 1999a; Smith et al. 2003).
Lumbsch and Huhndorf (2010) accepted only
Apiorhynchostoma, Aquasphaeria, Brunneiapiospora,
Clypeosphaeria, Crassoascus, Duradens, Palmomyces and
Pseudovalsaria in the family. Maharachchikumbura et al.
(2015) accepted Apioclypea, Aquasphaeria,
Brunneiapiospora, Clypeosphaeria, Crassoascus and
Palmomyces as the genera in Clypeosphaeriaceae, following
Lumbsch and Huhndorf (2010), as these genera, except for
Clypeosphaeria, lack sequence data.
Type: Clypeosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–
24: 117 (1870) [1869–70]
Fungal Diversity
Type species: Clypeosphaeria mamillana (Fr.) Lambotte,
Flore Mycologique Belge 2: 247 (1880)
Clypeosphaeria mamillana (Fr.) Lambotte, Flore
Mycologique Belge 2: 247 (1880)
Basionym: Clypeosphaeria notarisii Fuckel, Jb. nassau.
Ver. Naturk. 23–24: 117 (1870).
Facesoffungi number: FoF 00709; Fig. 9
Saprobic on decaying wood. Sexual morph: Clypeus
dome-shaped, well-developed, black. Ascomata 380–450μm
diam., 350–400μm high (x =428×389μm, n=10), immersed
under the clypeus, solitary, scattered, globose to subglobose,
coriaceous, black, ostiole central, papillate. Papilla short, narrow, internally lined with hyaline, filamentous periphyses.
Peridium 20–35μm wide (x =25μm, n=20), comprising outer, brown, elongated, flat, cells of textura angularis and inner,
hyaline, elongated, flat, cells of textura angularis.
Hamathecium comprising numerous, 3–5 μm wide (x =
3.5μm, n=20), hypha-like, aseptate, unbranched, flexuose,
paraphyses, tapering towards the apex. Asci 165–230×8–
11μm (x =198×10μm, n=20), 8-spored, unitunicate, cylindrical, short pedicellate, apex rounded, with 1.5–2.5μm diam.,
2.5–3.5μm high (x =2.1×2.8μm, n=20), wedge-shaped, J+
apical apparatus. Ascospores 20–25×6–9μm (x =20×7μm,
n=30), uniseriate, dark brown, ellipsoidal to fusiform, unicellular, straight or slightly curved, 0–5-distoseptate, not constricted at the septa, smooth-walled. Asexual morph:
Undetermined.
Material examined: GERMANY, Ahlen, Oestricher forest,
December 1823, K.W.G.L Fuckel (G00127177, holotype).
Notes: Clypeosphaeria is typified by Clypeosphaeria
mamillana (Fr.) Lambotte (Kang et al. 1999a). This genus
has been synonymized under several genera (Barr 1989),
while Hyde et al. (1998) treated Clypeosphaeria as a monotypic genus, excluding all the other species. However currently 41 species have been listed under genus Clypeosphaeria
(Mycobank 2015). Clypeosphaeria is characterized by 0–5pseudoseptate ascospores, without germ pores. The type species Clypeosphaeria mamillana is previously named as
Clypeosphaeria notarisii Fuckel (Kang et al. 1999a;
Mycobank 2015). However Clypeosphaeria notarisii was an
illegitimate name, because the name should be a combination
with Sphaeria clypeiformis De Not. (Mycobank 2015).
Discosiaceae Maharachch. & K.D. Hyde, fam. nov.
Index Fungorum number IF551177; Facesoffungi number:
0665
Saprobic or pathogenic on plant leaves. Sexual morph:
Ascomata scattered, immersed to erumpent. Ostiole circular,
papillate. Peridium comprising several layers of compressed
cells, brown at the base, dark brown outwardly. Hamathecium
comprising numerous, hypha-like, flexuose, paraphyses. Asci
8-spored, unitunicate, obclavate to cylindrical, with a discoid,
J+ subapical ring. Ascospores overlapping uniseriate or
biseriate, obovoid to ellipsoid, transversally septate, with or
Fig. 9 Clypeosphaeria mamillana (holotype). a Herbarium packet. b
Herbarium specimens. c Appearance of ascomata on host substrate. d
Cross section of ascoma. e Peridium. f, g J+ apical ring of ascus in
Melzer’s reagent. h–k Asci. l–o Ascospores. Scale bars: c=500 μm, d=
50 μm, e=10 μm, f, g=5 μm, h–k=10 μm, l–o=5 μm
without vertical septa, hyaline. Asexual morph:
Coelomycetous. Conidiomata stromatic, scattered to gregarious, subepidermal or subepidermal in origin, immersed to
erumpent. Peridium composed of 3–4 layers of cells of textura
angularis, outer 1–2-layers brown and inner 1–2 layers hyaline or pale brown. Conidiophores arising from the inner layer
cells of basal stroma, absent or reduced to conidiogenous
cells, when present long, septate and branched.
Conidiogenous cells holoblastic, annelidic, cylindrical or
lageniform, hyaline. Conidia cylindrical to clavate, or fusiform, straight or curved, 2–4-euseptate, hyaline, pale olivaceous, or brown, bearing cellular, filiform or attenuated
appendages.
Type: Discosia Lib., Pl. crypt. Arduenna, fasc. (Liège) 4:
no. 346 (1837)
Notes: Phylogenetic analyses of combined LSU and ITS
sequence data of Discosia and related genera show
Discosiaceae taxa to be distinct from the family
Bartaliniaceae and Pestalotiopsidaceae. Hence the new family Discosiaceae is introduced for Discosia and Adisciso,
Discostroma, Immersidiscosia, Sarcostroma, and
Seimatosporium (Fig. 1). Members of the family
Discosiaceae are saprobes, endophytes, or foliar pathogens
in tropical and temperate regions (Nag Raj 1993; Tanaka
et al. 2011). The family presently includes four coelomycetous
asexual genera Discosia, Immersidiscosia, Sarcostroma and
Seimatosporium, and two sexual genera Adisciso and
Discostroma; which were earlier classified under
Amphisphaeriaceae. Von Arx (1981) treated Bartalinia,
Bartaliniopsis, Bleptosporium, Doliomyces and Hyalotia as
synonyms of Seimatosporium. However, Nag Raj (1993) did
not accept von Arx’s (1981) treatment and sequence data also
does not support this.
Discosia Lib., Pl. crypt. Arduenna, fasc. (Liège) 4: no. 346
(1837)
Discosia was described by Libert (1837) with D. artocreas
(Tode) Fr. as the type. The coelomycetous Discosia is widespread in subarctic, temperate and tropical regions. Discosia
species are generally saprobes or plant pathogens and rarely
biotrophs (Crous et al. 2013).
Discosia neofraxinea W.J. Li, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF551176; Facesoffungi
number: FoF 00664; Fig. 10
Etymology: Named because its similarity with Discosia
fraxinea.
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Discosia neofagicola (holotype). a Herbarium specimen. b, c
Black conidiomata on the host. d Vertical section of conidioma. e, g
Section of peridium. h Germinating spore. f, i–l Conidiogenous cells
and developing conidia. m–o Conidia. Scale bars: b= 200 μm, c=
20 μm, d=100 μm, e, g=50 μm, h–j=10 μm, f, i–l=5 μm, m–n=5 μm
Fig. 10
Holotype: MFLU 15–0375
Saprobic on dead leaf of Fagus sylvatica. Sexual morph:
Undetermined. Asexual morph: Coelomycetous.
Conidiomata 125–176 μm high, 450–550 μm diam.,
stromatic, pycnidioid, solitary or gregarious, superficial, globose to subglobse, unilocular, rugose, dull black, with a short
beak, ostiolate. Ostiole 60–80μm diam., circular or oval,
opening to the exterior. Peridium 10–20μm wide, composed
of two types of pseudoparenchymatous cells, forming thickwalled cells of textura epidermoidea towards the apex, with
textura prismatica at the base, with inner layers hyaline gradually merging with the outer dark brown layers.
Conidiophores arising from the inner layer cells of the basal
stroma, septate, unbranched or occasionally branched, long,
cylindrical, hyaline, smooth. Conidiogenous cells 6–40μm
long × 1–2 μm wide, holoblastic, cylindrical, hyaline,
smooth-walled. Conidia 15–18×2.5–3.5μm (x =16×3μm;
n=30), median cells pale brown, naviculate to subcylindrical,
narrow towards the base, 3-septate, slightly curved, with cells
of unequal length, hyaline to pale brown, bearing appendages;
basal cell narrowly obconic with a truncate base bearing a
prominent dehiscence scar, with septa thicker and darker than
the periclinal wall, with the second cell close to apical cell
twice or more than two times that of the size of the third cell
close to basal cell, with apical cell subconical and with an
acute apex; appendage 10–20μm long, single, unbranched,
flexuous.
Materials examined: ITALY. Province of Forlì-Cesena
[FC], Santa Sofia, Campigna, on dead leaf of Fagus sylvatica,
24 June 2012, Erio Camporesi, IT 469 (MFLU 15–0375,
holotype); ex-type living culture, MFLUCC 13–0204,
ICMP; ibid. (HKAS 88184, isotype); ibid. Arezzo [AR],
Santa Sofia, near Passo della Braccina, on dead leaf of
Fagus sylvatica, 30 April 2013, Erio Camporesi, IT 469E
(MFLU 15–0736, paratype); living culture, MFLUCC 13–
0378; paratype); ibid. IT 469 F (MFLU 15–0737, paratype).
Notes: Discosia neofraxinea is similar to D. artocreas
(Tode) Fr. in having 3-septate, hyaline to pale brown conidia
with single appendages at each end, but differs in the form of
conidiomata and conidiogenous cells. Discosia neofraxinea
closely resembles D. fraxinea (Schwein.) Nag Raj which is
known from Fraxinus americana. Both share similarities in
the dimensions of conidia and conidiogenous cells, but differ
in the form of conidiomata (Nag Raj 1993). Discosia
neofraxinea has superficial, globose to subglobse, unilocular,
rugose, conidiomata, which are much smaller than the
applanate to discoid, erumpent, unilocular to bilocular,
glabrous, occasionally rugose conidiomata in D. fraxinea
(700–1000μm diam., up to 80μm high). Phylogenetically,
the strains of D. neofraxinea formed a discreet branch in
Discosia. Based on a combination of morphology and phylogeny, we introduce D. neofraxinea as a new taxon.
Discosia pseudopleurochaeta Wijayawardene, Camporesi
& K.D. Hyde, sp. nov.
Index Fungorum number: IF551179; Facesoffungi
number: FoF 00661; Fig. 11
Etymology: Named after its morphological similarity with
Discosia pleurochaeta
Holotype: MFLU 15–0743
Saprobic on dead leaf of Crataegus sp. Sexual morph:
Undetermined. Asexual morph: Coelomycetous.
Conidiomata 300–360μm diam., 40–80μm high, stromatic,
gregarious, occasionally confluent, flattened, glabrous, black,
superficial, unilocular. Ostiole papillate, single circular, central. Conidiomata wall thick at the base, 15–25μm, composed
of pale brown, thin-walled cells of textura angularis, wall near
ostiole thinner, composed of dark brown, thick-walled cells of
textura angularis. Conidiogenous cells 3–5×1–3μm, holoblastic, discrete, determinate, conical, hyaline to pale brown,
smooth, only in basal conidiomatal wall. Conidia 20–28×3–
5μm (x =23.6×4.11μm, n=20), median cells pale brown,
apical and basal cells sub-hyaline to hyaline, subcylindrical,
tapered to a truncate base, apex obtuse, straight or slightly
curved, dorsiventral, with 3-transverse septate, smoothwalled, bearing appendages at both ends; apical appendage
15–18μm, long, tubular, unbranched; basal appendage 17–
22μm, long, tubular, unbranched.
Materials examined: ITALY, Province of Forlì-Cesena
[FC], Collinaccia - Castrocaro e Terra del Sol, dead and hanging leaf of Crataegus sp, 20 October 2012, Erio Camporesi, IT
836 (MFLU 15–0743, holotype); ibid (HGUP530 paratype)
Notes: Farr and Rossman (2015) list Discosia artocreas
(Tode) Fr. and Discosia faginea Lib. from Crataegus spp.
Nag Raj (1993) listed the latter species under ‘excluded or
unexamined taxa’ as its type species was not available.
Discosia artocreas has smaller conidia (conidial dimensions
14–22×2–3μm) than our collection and its appendages are
inserted about 2–3 μm from the apical and basal septum
(Nag Raj 1993), but in our collection both apendages are
inserted very close (1–2μm) to the apical and basal septa.
Further we checked Nag Raj (1993) to find other species
which have close morphological characters with our collection. Discosia pleurochaeta Mont. & Durieu (19.5–27×3–
3.5μm) has similar conidial dimensions, thus resembling our
collection, but has narrower conidia. Further, D. pleurochaeta
has longer conidiogenous cells (7–11×2.5–3μm) than our collection. Hence, we introduce a new species to accommodate
our new collection based on morphology.
Seimatosporium Corda, Deutschl. Fl., 3 Abt. (Pilze
Deutschl.) 3 (13): 79 (1833)
Fungal Diversity
Facesoffungi number: FoF 00842
Saprobic on leaves, stem and bark of various plants,
pathogenic on leaves of Eucalyptus spp. Sexual morph:
Discostroma-like. Asexual morph: Conidiomata acervular,
solitary to gregarious, superficial to immersed, unilocular, globose to subglobose, dark brown to black. Conidiomata wall
composed of brown thin or thick-walled textura angularis.
Conidiophores cylindrical, filiform, septate, branched, hyaline. Conidiogenous cells holoblastic, annellidic, integrated or discrete, hyaline, determinate. Conidia variable, cylindrical, fusiform or clavate or obovoid, (2
−)3(−5)-septate, aguttulate, medium cells brown, basal
cell hyaline, with or without basal appendage, or with
the apical cell provided with a single, cellular, simple or
branched appendage, basal cell truncate, continuous or
occasionally constricted at the septa.
Fig. 12 Discostroma fuscellum (MFLU 15–0750). a–c Ascomata on
host surface. d, e Section of ascoma. f Peridium. g Paraphyses. h–j
Immature asci. k Mature ascus l Mature ascus in Congo red m–o
Immature ascospores. p–q Mature ascospores. r Ring-like apical
apparatus of ascus in Congo red s Germinating ascospore. t, u Culture
on MEA. Scale bars: a–b, h–l=200 μm, c–e=100 μm, f–g, r=50 μm, m–
q=20 μm, s=5 μm, t–u=30 mm
Notes: Corda (1833) introduced Seimatosporium based on
S. rosae as the type species. Sutton (1963), Shoemaker and
Muller (1964) and Nag Raj (1993) provided taxonomic notes
on the genus. Von Arx (1981) treated Bartalinia,
Bartiloniopsis, Bleptosporium, Doliomyces and Hyalotia as
generic synonyms of Seimatosporium. Nag Raj (1993) however, accepted Seimatosporium as a valid genus.
Nag Raj (1993) mentioned Seimatosporium has
Discostroma sexual morphs. Okane et al. (1996) and
Fig. 11 Discosia pseudopleurochaeta (holotype). a Conidiomata on leaf. b, c Cross section of stromata. d Cross section of stromatic wall. e–h
Developing conidia attached to conidiogenous cells. i, j Conidia. Scale bars: b, c=50 μm, d=25 μm, e–h=8 μm, i, j=20 μm
Fungal Diversity
Fungal Diversity
Hata key ama an d H ar ada (2 004 ) al s o in t r od uce d
Seimatosporium spp. with Discostroma sexual morphs.
Barber et al. (2011) and Tanaka et al. (2011) also showed that
both Seimatosporium and Discostroma cluster in a monophyletic clade in their molecular data analyses. However, both
genera lack molecular data for their type species (Tanaka
et al. 2011), thus we prefer to maintain both names until the
link is proven by molecular analyses.
Discostroma Clem., Gen. fung. (Minneapolis): 50, 173
(1909)
Notes: Species of Discostroma and Seimatosporium have
been linked by molecular data (Tanaka et al. 2011), however
the molecular links have not been proven for the types of
either genera. Both names should therefore be retained pending fresh collections of the types and a molecular study.
Discostroma fuscellum (Berk. & Broome) Huhndorf [as
‘fuscella’], Bull. Ill. nat. Hist. Surv. 34 (5): 520 (1992)
Index Fungorum number: IF 359431; Facesoffungi
number: FoF 00697; Fig. 12
Saprobic on dead stem of Rosa canina, forming conspicuous, rounded, brown to black ascomata. Sexual morphs:
Ascomata 280–478μm diam., 149–227μm high (x =379×
188 μm, n = 30), solitary or clustered, immersed, semiimmersed or superficial, globose to subglobose, membranous,
brown, ostiolate. Peridium 25–44μm wide (x =35μm, n=30),
three-layered, outer layer comprising brown, thick-walled
cells of textura angularis, central layer comprising light
brown to yellow, thin-walled cells of textura angularis, inner
layer comprising hyaline, thin-walled cells of textura
angularis. Hamathecium comprising 3–6μm wide (x =4μm,
n=30), numerous, hypha-like, septate, paraphyses, tapering
distally, not embedded in a gelatinous matrix. Asci 103–
127×8–11μm (x =115×10μm, n=30), 8-spored, unitunicate,
cylindrical, with pedicel often tapering to a point, apically
rounded or truncate, with a refractive, J+, bipartite, apical ring.
Ascospores 15–18×6–8μm (x =17×7μm, n=30), uniseriate
or overlapping uniseriate, fusiform or ellipsoidal, 0–4-septate,
hyaline, smooth-walled, thick-walled. Asexual morph:
Undetermined.
Culture characters: Colonies growing on PDA, reaching
7 cm diam. after 3 weeks at 20–25 °C, with whitened, dense,
aerial mycelium on the surface, reverse similar in colour.
Material examined: ITALY. Province of Forlì-Cesena [FC],
Poggio Baldo - Predappio Alta, on dead stem of Rosa canina,
3 March 2013, Erio Camporesi, IT 1093 (MFLU 15–0750,
MFLU 15–0751, MFLU 15–1134, MFLU 15–1135); living
culture, MFLUCC 14–0052, ICMP.
Notes: There are 30 species recorded under Discostroma
(Index Fungorum, 2015) and Paulus et al. (2006) provided a
key to 28 Discostroma species. We consider our species to be
Discostroma fuscellum based on morphology. Discostroma
fuscellum was stated to have immersed perithecia with short
protruding papillae, cylindrical asci, with a J+ apical ring and
Fig. 13 Seimatosporium cornii (holotype) a, b Conidiomata on Cornus
sp. c, d Cross sections of conidiomata. e Paraphyses. f–k Developing
conidia attach to conidiogenous cells. l–n Conidia. Scale bars: c,
d=100 μm, e=30 μm, f–n=15 μm
hyaline, 3-septate, elliptical, pigmented ascospores (Paulus
et al. 2006). In our collection the ascospores were hyaline
which agrees with Huhndorf (1992). Eriksson (2009) used
Leptosphaeria fuscella for Discostroma fuscellum, however,
the morphology and phylogenic analyses, show it to belong to
Discostroma and is close to Seimatosporium and Tanaka et al.
(2011) proposed to use Discostroma fuscellum for this taxon.
Seimatosporium cornii Wijayawardene, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF551180; Facesoffungi
number: FOF 00662; Fig. 13
Etymology: Named after the host genus on which the fungus occurs.
Holotype: MFLU 15–0742
Saprobic on dead branches of Cornus sp. Sexual morph:
Undetermined. Asexual morph: Conidiomata 300–330μm
diam., 150–175μm high, acervular, unilocular, subglobose,
superficial to subepidermal, solitary, dark brown, apapillate
ostiolate. Conidiomata wall multi-layered, outer wall thick,
composed of brown cells of textura angularis, inner wall thin,
hyaline. Paraphyses 30–60μm, cylindrical, filiform, hyaline,
aseptate. Conidiophores 13–30×2–4μm, long, cylindrical,
branched, hyaline, smooth-walled. Conidiogenous cells holoblastic, annelledic, simple, integrated, determinate, hyaline.
Conidia 21–29×9–11 (x =24.9×9.93, n=20), median cells
brown to dark brown, basal and apical cells subhyaline
to pale brown, clavate or obovoid, occasionally truncate
at base, straight to slightly curved, with 3-transverse
septate, dark brown septa, aguttulate, smooth-walled, appendage absent.
Culture characteristics: slow growing, reaching 2.5–3 cm
in 7 days at 18 °C, on PDA, white from above, grey from
below, circular, with uneven margin, not zonate, with thin
mycelium, becoming dense after 14 days
Materials examined: ITALY, Province of Pesaro-Urbino
[PU], Monte Nerone, on branches of Cornus sp., 11
June 2012, Erio Camporesi, IT 426 (MFLU 15–0742,
holotype); ibid (HGUP531 paratype); ex-type living culture,
MFLUCC 14–0467, GUCC 0058.
Notes: Farr and Rossman (2015) reported Seimatosporium
lichenicola (Corda) Shoemaker & E. Müll. (Conidial dimension 13–15 × 5.5–6.5 μm fide Sutton 1980) and
Seimatosporium salicinum (Corda) Nag Raj (11–17×4–6μm
fide Nag Raj 1993) from Cornus spp. However, our collection
is morphologically distinct from these two species and other
known species in Sutton (1980) and Nag Raj (1993). Hence,
we introduce this new species based on morphology and molecular data analyses.
Fungal Diversity
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Seimatosporium rhombisporum Senan. Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF551195; Facesoffungi
number: FoF 00693; Fig. 14
Etymology: Species name refers to the rhomboid conidia.
Holotype: MFLU 15–0727
Saprobic on dead branches of Vaccinium myrtillus L.
Sexual mor ph: Unde termined. Asexual morph:
Conidiomata 65–75μm high, 105–125μm diam., (x =70×
120 μm, n = 20), acervular, uniloculate, subglobose to
Fig. 14 Seimatosporium rhombisporum (holotype). a Vaccinium
myrtillus stems with conidiomata. b, c Conidiomata. d, e Cross section
of the conidiomata. f–j Conidia attached to conidiogenesis cells and
conidiophores. k–o Conidia. p Germinating conidia. Scale bars: b=
200 μm, c=500 μm, d=50 μm, e–p=10 μm
Fungal Diversity
subconical, erumpent to superficial, solitary, dark brown.
Peridium multi-layered, comprising brown, cells of textura
angularis to textura globosa. Conidiophores 1–1.5μm high,
0.5–1μm wide (x =1.3×0.7μm, n=20), cylindrical, branched,
hyaline, smooth-walled. Conidiogenous cells 5.5–12.5μm
high, 0.5–1μm diam., (x =7.5×0.7μm, n=20), cylindrical,
unbranched, holoblastic, anneledic, simple, integrated, hyaline. Conidia 10.5–12×3–4μm (x =11×3.6μm, n=20), clavate, with brown to dark brown median cells and hyaline to
pale brown basal and apical cells, pointed at apex, blunt at
base, straight to slightly curved, 3-transverse septate, septa
thick, smooth-walled, lacking appendages with germinating
hyphae originating from basal hyaline cell during
germination.
Culture characters: Growing on MEA, reaching 1 cm
within 7 days when incubated at 18 °C, flat, circular, white
colony with smooth margin.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Santa Sofia, Monte Falco, on Vaccinium myrtillus
(Ericaceae), 7 May 2013, Erio Camporesi IT 1228,
(MFLU15–0727, holotype) ibid., (MFLU15–0727bis,
paratype); living culture MFLUCC 15–0543, ICMP.
Notes: We introduce Seimatosporium rhombisporum sp.
nov. which differs from other species in having erumpent,
conidiomata with long, immersed conidiogenous cells.
Conidiomata are filled with conidiogenous cells and conidia
form at the apex. The dark, rhomboid conidia are quite specific and phylogenetic analysis (Fig. 1) indicates the uniqueness of this species. However, Seimatosporium vaccinii
(Fuckel) B. Erikss. also has conidia without appendages and
was obtained from the same host (Lee et al. 2006). The colour
and shape of conidia are, however, different. Based on a
megablast search of GenBank, the closest hit using LSU sequence data to our strain are Sarcostroma bisetulatum (Guba)
Nag Raj (CBS 122695, EU552155; Identities = 883/897
(98 %), Gaps=8/897 (0 %)), Seimatosporium obtusum (H.J.
Swart & M.A. Will.) P.A. Barber & Crous (CPC 12935,
JN871215; Identities=876/893 (98 %), Gaps=7/893 (0 %)),
and S. eucalypti (McAlpine) H.J. Swart (CPC 158, JN871211;
Identities=874/892 (98 %), Gaps=8/892 (0 %)). Sarcostroma
was introduced to accommodate species having 2–5-septate
conidia with only a basal appendage or without any
appendages and Sutton (1980) synonymised Sarcostroma under Seimatosporium. Both Seimatosporium obtusum and
Seimatosporium eucalypti (McAlpine) H.J. Swart have long,
falcate, fusiform or sigmoid conidia with 3–4-septa, rarely 5septa, and basal appendages. These species were excluded
from Seimatosporium and included in Vermisporium based
on their morphology and phylogeny (Barber et al. 2011).
Therefore we introduce Seimatosporium rhombisporum as a
new species considering both morphology and phylogeny.
Seimatosporium ficeae Dayarathne, Phookamsak & K.D.
Hyde, sp. nov.
Index Fungorum number: IF551175; Facesoffungi
number: FOF 00633; Fig. 15
Etymology: Name reflects the host Ficus, from which this
species was isolated.
Holotype: MFLU 15–0703
Saprobic on living leaf of Ficus sp. Sexual morph:
Undetermined. Asexual morph: Conidiomata 50–60 μm
high, 45–90μm diam., acervuli, globose, submerged, solitary,
black. Peridium 10–15μm wide, composed of hyaline to pale
brown cells of textura angularis to textura globosa.
Conidiophores up to 10 μm long, subcylindrical.
Conidiogenous cells 13–22μm long, holoblastic, cylindrical
to ampulliform, hyaline, thin-walled, smooth, arising from the
basal layers of conidiomata. Conidia 15–21×5.5–7.5μm (x =
18×6.5μm, n=30), fusiform, straight, 5-septate, concentric at
the septa, with 4 median cells, brown, terminal cells, hyaline,
bearing single 8–20μm long, unbranched, central appendages
at both ends.
Culture characters: Colonies on PDA slow growing,
reaching 25 mm diam. after 2 weeks at 25–30 °C, upper side
circular, white to gray, and cottony, lower side circular, white
to gray.
Material examined: CHINA, Yunnan Province, ShangriLa, on leaf spots of Ficus sp., 30 October 2014, R.
Phookamsak SGL 002 (MFLU 15–0703 holotype); ex-type
living culture, MFLUCC 15–0519=ICMP.
Notes: The main differences between Seimatosporium
ficeae and other species in Seimatosporium lie in the morphology of the conidia. The conidia of S. ficeae are 5-septate conidia with four, brown median cells and hyaline terminal cells
at both ends. S. consocium (Peck) Shoemaker also posses
similar morphological characters to S. ficeae, including 5septate with four brown to light brown median cells.
Meanwhile, the conidial septation of S. hysterioides (Fuckel)
Brockmann also exhibits a similar pattern with five septa but,
abundantly they are 3–4-septate. Both S. consocium and
S. hysterioides bear central apical appendages and eccentric
basal appendages on conidia whereas S. ficeae comprises of
central appendages at both ends. In the phylogenetic analysis
(Fig. 1), S. ficeae is related to S. hypericinum (Ces.) B. Sutton
which has curved, falcate, 4-celled conidia. However, they are
well-separated with a high bootstrap support.
Seimatosporium vitis Y.P. Xiao, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF551200; Facesoffungi
number: FOF 00698; Fig. 16
Etymology: Name referring to the host genus on which the
species occurs Holotype: MFLU 15–0746
Saprobic on dead stems of Vitis vinifera, forming conspicuous, rounded, black conidiomata. Sexual morph:
Undetermined. Asexual morph: Coelomycetous.
Conidiomata 274–368 μm diam., 112–162 μm high (x =
321×137μm, n=30), acervuli, solitary or gregarious, semi-
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immersed, dark brown to black, elongate to rounded or irregular in outline. Peridium 25–39μm wide (x =32μm, n=30),
thin, comprising light brown, thin-walled cells of textura
angularis. Conidiophore reduced to conidiogenous cells.
Conidiogenous cells 1–2μm diam., 13–36μm long (x =1×
25μm, n=30), holoblastic, cylindrical, hyaline to pale brown,
thick-walled, smooth. Conidia 34–40×14–17μm (x =15×
37 μm, n = 60), ellipsoid or fusiform, straight or slightly
curved, 3-euseptate, constricted at septate, guttulate, bearing
an appendage at the base, apical cell conical with broadly
rounded apex, two median cells sub-cylindrical, brown to dark
brown, relatively thick-walled than the end cells, basal cell
Fig. 16 Seimatosporium vitis (holotype) a, b Conidiomata on host
surface. c Vertical section of conidioma. d–f Base of conidioma with
conidiogenous cells, conidiophores and developing conidia. g–k Mature
conidia. l Germinating conidium. m Culture from above on MEA
medium. n Culture from below on MEA medium. Scale bars: a=
2.5 mm, b=200 μm, c–d=100 μm, e=40 μm, f–k=20 μm, l=10 μm,
m–n=30 mm
obconical, pale brown, bearing appendages. Appendages 4–
8μm long (x =5μm, n=30), central or eccentric, single, unbranched, filiform.
Culture characters: growing on PDA, reaching 7 cm diam.,
after 2 weeks at 20–25 °C, with circular, whitened to brown,
Fig. 15 Seimatosporium ficeae (holotype). a Pycnidium on host b, c Conidiomata on host tissue. d Section through conidiomata. e Section through
peridium. f Conidiogenous cells. g–j. Conidia. k–l Culture of Seimatosporium sp. on PDA. Scale bars: d–e=100 μm, f=50 μm, g–j=10 μm
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dense, aerial mycelium on the surface, reverse similar in
colour.
Material examined: ITALY. Province of Forlì-Cesena [FC],
near Predappio Alta, on dead stem of Vitis vinifera L.
(Fabaceae), 3 March 2013, Nello Camporesi, IT1092
(MFLU 15–0746, holotype); IT 1092 (MFLU 15–0747,
MFLU 15–0748, 15–0749, paratypes); ex-type living cultures, MFLUCC 14–0052, ICMP, MFLUCC 14–0051.
Notes: Seimatosporium vitis is similar to S. pistaciae Crous
& Mirab. which has conidia with an obtusely rounded apical
cell, with a single apical appendage (Crous et al. 2014b).
Seimatosporium vitis however, differs in having 3-septa, a
conical, apical cell with broadly rounded apex, brown to dark
brown, median cells and a pale brown basal cell bearing an
appendage. This is the first Seimatosporium species to be described from Vitis.
Pestalotiopsidaceae Maharachch. & K.D. Hyde, fam. nov.
Index Fungorum number IF551178; Facesoffungi number:
0666
Saprobic or pathogenic on plants. Sexual morph:
Ascomata scattered, solitary or in small group, immersed,
apex short erumpent. Ostiole subglobose, papillate ostiolar
canal periphysate. Peridium comprising several layers of dark
brown to blackish, compressed cells. Hamathecium comprising tapering paraphyses. Asci 8-spored, unitunicate, cylindrical, short to long pedicellate, with a J+ or J- apical ring.
Ascospores uniseriate, brown, ovoid to elliptic, straight or
inequilateral, 2–3-septate. Asexual morph: Coelomycetous.
Conidiomata acervular or pycnidial, subglobose, globose, clavate, solitary or aggregated, dark brown to black, immersed to
erumpent, unilocular or irregularly plurilocular. Peridium
comprising 2–3 strata of textura angularis, outer layer of pale
brown, thick-walled cells, becoming hyaline in the inner layer.
Conidiophores indistinct, often reduced to conidiogenous
cells, when present peripheral, hyaline, branched or unbranched, cylindrical or lageniform. Conidiogenous cells discrete, holoblastic, annellidic, indeterminate, integrated, cylindrical, hyaline, smooth. Conidia ellipsoid to clavate, or fusiform, straight or curved, 3–4-euseptate, hyaline, pale olivaceous or brown, bearing cellular, filiform or attenuated
appendages.
Type: Pestalotiopsis Steyaert, Bull. Jard. bot. État Brux. 19:
300 (1949)
Notes: The new family Pestalotiopsidaceae, which possesses pestalotiopsis-like asexual morphs is introduced for
Ciliochorella, Lepteutypa, Monochaetia, Neopestalotiopsis,
Pestalotiopsis, Pseudopestalotiopsis and Seiridium (Fig. 1).
Maharachchikumbura et al. (2014) suggested that the monotypic genus Pestalotia (1839) might be a synonym of
Seiridium (1816), since both genera have similar morphologies. Members of the Pestalotiopsidaceae are common phytopathogens that cause a variety of diseases in plants or are
often isolated as saprobes or endophytes and are widely
distributed throughout tropical and temperate regions (Guba
1961; Barr 1975; Nag Raj 1993; Maharachchikumbura et al.
2014). The sexual morph of Pestalotiopsis is Pestalosphaeria
(Barr 1975) and Seiridium has a Lepteutypa sexual morph
(Nag Raj and Kendrick 1985).
Pestalotiopsis Steyaert, Bull. Jard. bot. État Brux. 19: 300
(1949)
Notes: Based on the conidial forms, Steyaert (1949) split
Pestalotia into three genera, namely Pestalotia, Pestalotiopsis
and Truncatella. Pestalotiopsis was introduced for species
with 5-celled conidia and P. maculans is regarded as the type
species. The conspicuous character of this genus is 5-celled,
fusiform conidia, with three-coloured median cells, hyaline
end cells, and one or more apical appendages. The sexual
morph of Pestalotiopsis is Pestalosphaeria and only 13 species are known as compared to the asexual morph (254 species
names). Maharachchikumbura et al. (2011) pointed out that
the common Pestalotiopsis name should be applied to both
morphs.
Neopestalotiopsis Maharachch. et al., Stud. Mycol. 79: 135
(2014)
Notes: Maharachchikumbura et al. (2014) resolved genera
in the Amphisphaeriaceae based on analysis of LSU sequence
data. Besides accepting Pestalotiopsis, they introduced
Neopestalotiopsis and Pseudopestalotiopsis.
Morphologically Neopestalotiopsis can easily be distinguished from Pseudopestalotiopsis and Pestalotiopsis by its
versicolorous median cells. Furthermore, in
Neopestalotiopsis, conidiophores are indistinct and often reduced to conidiogenous cells (Maharachchikumbura et al.
2014).
Pseudopestalotiopsis Maharachch. et al., Stud. Mycol. 79:
180 (2014)
Notes: Pseudopestalotiopsis was segregated from
Pestalotiopsis based on the type Pseudopestalotiopsis theae
(Sawada) Maharachch. et al. (Maharachchikumbura et al.
2014). They designated an epitype for Pestalotiopsis theae
(Sawada) Steyaert from fresh leaves of Camellia sinensis collected in Thailand (Maharachchikumbura et al. 2013).
Pseu do pes talo tiop sis c a n be di s t i n g u i s h e d f r o m
Neopestalotiopsis and Pestalotiopsis by sequence data and
in having generally dark, concolourous, median cells with
indistinct conidiophores (Maharachchikumbura et al. 2014).
Monochaetia (Sacc.) Allesch., Rabenh. Krypt.-Fl., Edn 2
(Leipzig) 1 (7): 665 (1902) [1903]
Notes: The genus Monochaetia was introduced by
Allescher (1902), with M. monochaeta (Desm.) Allesch. as
its type. No sexual morphs are conclusively known for this
genus. Steyaert (1949) did not retain Monochaetia as a distinct
genus, and transferred many of the species of Monochaetia
into Pestalotiopsis or Truncatella. However, Monochaetia
was considered as a distinct genus by Guba (1961) based on
its single apical appendage. LSU phylogenetic analyses reveal
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Monochaetia to represent a distinct genus from Pestalotiopsis,
Seiridium and Truncatella (Maharachchikumbura et al. 2014).
It is essential to incorporate molecular data and more taxon
sampling in future analyses as Monochaetia includes 3-, 4-,
and 6-celled conidial forms (Maharachchikumbura et al.
2014).
Seiridium Nees, Syst. Pilze (Würzburg): 22 (1816) [1816–
17]
Notes: Sequence data has revealed Seiridium to represent a
distinct genus in the family Amphisphaeriaceae, which is
characterised by 6-celled conidia (Jeewon et al. 2003b;
Maharachchikumbura et al. 2014) and in the present study it
is placed in Pestalotiopsidaceae. Blogiascospora and
Lepteutypa have been identified as the sexual morph of
Seiridium. However, Seiridium is the oldest and most commonly used name and therefore, should be applied to represent
both morphs.
Ciliochorella Syd., in Sydow & Mitter, Annls mycol. 33
(1/2): 62 (1935)
Notes: The genus Ciliochorella was introduced by Sydow
and Mitter (1935), with C. mangiferae Syd. as its type. The
sexual morph of the genus has not been reported (Hyde et al.
2011), and the characteristic feature of the genus is
eustromatic, pycnidial conidiomata; indistinct conidiophores;
enteroblastic or phialidic conidiogenesis and 3-euseptate conidia with pale brown median cells and cellular appendages
(Nag Raj 1993). Presently seven species are known in
Ciliochorella Tangthirasunun et al. (2015).
Phlogicylindriaceae Senan. & K.D. Hyde, fam. nov.
Index Fungorum Number: IF551190; Facesoffungi
number: FoF 00681.
Saprobic on leaves, twigs and branches, presently only
known from dicotyledons. Sexual morph: Pseudostroma
scattered, solitary, immersed, black, thick around papilla, thinner towards the base. Ascomata forming under pseudostroma,
solitary, globose, coriaceous, black, ostiolate, papillate.
Papilla short, not prominent. Peridium comprising outer,
brown, thick-walled cells of textura angularis and inner, hyaline to light brown, thin-walled cells of textura angularis.
Hamathecium comprising relatively wide, septate, hyaline
paraphyses. Asci 8-spored, unitunicate, cylindrical, with
knob-like pedicel, apically rounded, with J+, discoid
subapical ring. Ascospores overlapping uniseriate, hyaline, fusiform, equally 1-septate. Asexual morph:
Coelomycetous Conidiomata slimy, erect tufts of hyaline conidial masses, synnematous, indeterminate, turning brown with age. Conidiophores consisting of an
intricate network of brown, smooth, branched cells.
Conidiogenous cells hyaline, annellidic, smooth, becoming pale brown with age, ampulliform with elongated
necks, with percurrent proliferations. Conidia formed
apically, hyaline, cylindrical with obtusely rounded
apex, aseptate to 1-septate.
Notes: The new family Phlogicylindriaceae is introduced
to accommodate the genus Phlogicylindrium Crous et al.
(Summerell et al. 2006). The type species, Phlogicylindrium
eucalypti Crous et al. has no known sexual morph and occurred on lesions of living leaves of Eucalyptus in association
with a Mycosphaerella species. Phlogicylindrium is unique in
having slimy conidiomata, with erect tufts of hyaline conidial
masses, and hyaline, unicellular to 1-septate, cylindrical conidia, with obtusely rounded apices (Summerell et al. 2006).
The genus comprises three species (Index Fungorum 2015).
During studies on Italian ascomycetes, we collected two species that cluster with Phlogicylindrium. Thus we introduce a
second genus to Phlogicylindriaceae for which we know the
sexual characters.
Type: Phlogicylindrium Crous et al., in Summerell et al.,
Fungal Diversity 23: 340 (2006)
Type species: Phlogicylindrium eucalypti Crous et al., in
Summerell et al., Fungal Diversity 23: 340 (2006); Fig. 17.
Ciferriascosea Senan., Bhat, Camporesi & K. D. Hyde,
gen. nov.
Index Fungorum Number: IF551191; Facesoffungi
number: FoF 00682.
Etymology: in honour of late Dr Raffaele Ciferri, an eminent mycologist from Italy.
Saprobic on dead plant matter. Sexual morph: Appearing
as scattered, black spots, slightly raised on the host surface.
Pseudostromata scattered, solitary, immersed, black.
Ascomata scattered, solitary, globose, coriaceous, brown to
black, ostiolate, papillate. Ostiolar papilla short, narrow, inner
wall lined by hyaline periphyses. Peridium comprising outer,
brown, thick-walled, layer of cells of textura angularis and
inner, hyaline, thick-walled, layer of cells of textura angularis.
Hamathecium comprising filamentous, septate, branched, hyaline paraphyses. Asci 8-spored, unitunicate, cylindrical, short
pedicellate, with an indistinct, J+ apical apparatus. Ascospores
overlapping uniseriate, hyaline, fusiform with narrowly
rounded, ends, median uniseptate, slightly constricted at the
septum. Asexual morph: Undetermined.
Notes: Ciferriascosea is introduced to accommodate two
taxa from Spartium junceum , collected in Italy.
Morphologically these two species are almost similar.
However they differ in dimension of the ascomata, asci and
ascospores dimensions. The most obvious character to differentiate these s p e c i e s a r e t h e a s c o s p o r e s e p t a ;
C. rectamurum has straight, thick septa while
C. fluctamurum has irregular, thin septa. Here we introduced this genus with C. rectamurum as the type species. However we could not obtain the asexual morph
from the culture. Combined gene analysis of LSU and
ITS sequences of this study shows the close affinity of
Ciferriascosea species to Phlogicylindrium species.
Ciferriascosea rectamurum Senan., Bhat, Camporesi &
K.D. Hyde, sp. nov.
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Index Fungorum Number: IF551193; Facesoffungi
number: FoF 00683; Fig. 18
Etymology: Species name based is on the straight, even
septa of the ascospores.
Holotype: MFLU 15–0726
Saprobic on dead stem of Spartium junceum L. appearing
as clear, scattered, black spots, slightly raised on the host surface. Sexual morph: Pseudostromata scattered, solitary, immersed, black. Ascomata 250–360μm high, 340–400μm
diam., (x = 265 × 370 μm, n = 20), scattered, solitary,
subglobose, coriaceous, black, ostiolate, papillate. Ostiolar
papilla short, not prominent, internally covered by periphyses.
Peridium 10–20μm wide, (x =15μm, n=20), comprising 4–6
outer layers of brown, thick-walled, cells of textura angularis
and 3–5 inner layers of hyaline, thick-walled cells of textura
angularis. Hamathecium comprising 2.5–3.5μm wide (x =
2.4μm, n=20), filamentous, septate, branched, paraphyses.
Asci 120–140×10–13μm (x =127×12μm, n=20), 8-spored,
unitunicate, cylindrical, pedicellate, apically rounded, apical
apparatus J+, not prominent. Ascospores 20–26×6–8μm (x =
23×7μm, n=20), overlapping uniseriate, hyaline, fusiform,
upper end pointed, lower end rounded, slightly curved,
Fig. 18 Ciferriascosea rectamurum (holotype). a Appearance of taxon
on host substrate. b. Cross section of pseudostroma and ascoma. c
Peridium. d–f Asci. g Paraphyses. h J+ apical ring of ascus mounted in
Melzer’s reagent. i–l Ascospores. m Germinating ascospore. n Upper
surface of culture. o Lower surface of culture. Scale bars: a=1 mm, b=
100 μm, c–f, h=20 μm, g, i–m=10 μm
equally uniseptate, septum straight, slightly constricted at the
septum, smooth-walled. Asexual morph: Undetermined.
Culture characters: Cultures growing on MEA reaching
1 cm within 7 days when incubated at 18 °C, circular, slightly
raised, smooth, entire margin, white, and woolly, with aerial
mycelium.
Materials examined: ITALY, Province of Forlì-Cesena,
Santa Sofia, Castellaccio di Corniolino, on branches of
Spartium junceum L (Fabaceae), 29 December 2012, Erio
Camporesi, IT 986 (MFLU 15–0726, holotype), ibid.,
(MFLU 15–0726Q, MFLU 15–0726R, paratypes); ex-type
living cultures MFLUCC 15–0542, ICMP.
Ciferriascosea fluctamurum Senan., Bhat, Camporesi &
K. D. Hyde, sp.nov.
Index Fungorum number: IF551194; Facesoffungi
number: FoF 00684; Fig. 19.
Fig. 17 Phlogicylindrium eucalypti (CBS H-19749, illustration based on Summerell et al. 2006). a Colony sporulating on MEA. b–d Conidiophores
and conidiogenous cells in vitro. e Conidiogenous cells. f Conidia. Scale bars: 10 μm
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Etymology: species name based on irregular septa in
ascospores.
Holotype: MFLU 15–0725
Saprobic on dead stem of Spartium junceum L., appearing
as slightly raised, scattered, black spots, on the host surface.
Sexual morph: Pseudostromata scattered, solitary, immersed,
black, clypeate. Ascomata 140–190μm high, 115–170μm
diam., (x =176×125μm, n=20), scattered, solitary, globose,
brown, ostiolate, papillate. Ostiolar papilla short, distinct, narrow. Peridium 5–8μm wide (x =6.2μm, n=20), comprising
4–6 outer layers of brown, thick-walled, cells of textura
angularis and 3–5 inner layers of hyaline, thick-walled cells
of textura angularis. Hamathecium comprising 1.2–2.3μm
wide (x =1.8μm, n=20), filamentous, septate, branched, paraphyses. Asci 60–75×6.5–8μm (x =66×7.3μm, n=20), 8spored, unitunicate, cylindrical, apex rounded, base with short
pedicel, apical apparatus J+, not prominent. Ascospores 9–
10×3–4μm (x =9.4×3.5μm, n=20), overlapping uniseriate,
hyaline, fusiform, with narrowly rounded ends, median
uniseptate, with a irregular septum, slightly constricted at the
septum. Asexual morph: Undetermined.
Culture characters: Cultures growing on MEA reaching
1 cm within 7 days when incubated at 18 °C, circular, slightly
raised, adpressed to the substrate, smooth, entire margined,
white, with sparse aerial mycelium.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Santa Sofia, Castellaccio di Corniolino, on branch of Spartium
junceum L (Fabaceae), 24 March 2013, Erio Camporesi, IT
1141 (MFLU 15–0725, holotype), (MFLU 15–0725bis,
isotype); ex-type living cultures MFLUCC 15–0541, ICMP.
Notes: Ciferriascosea fluctamurum occurs on Spartium
junceum and differs from Ciferriascosea rectamurum in having smaller asci and ascospores.
Xylariales
The order Xylariales is well-recognized
(Maharachchikumbura et al. 2015) and treated with 11 families in this paper. Xylariales comprises Apiosporaceae,
Cainiaceae, Coniocessiaceae, Diatrypaceae,
Graphostromataceae, Hyponectriaceae, Iodosphaeriaceae,
Lopadostomaceae fam. nov., Pseudomassariaceae fam.
nov., Vialaeaceae and Xylariaceae.
Apiosporaceae K.D. Hyde et al., in Hyde et al., Sydowia
50 (1): 23 (1998)
Facesoffungi number: FoF 00629
Saprobic or pathogenic on leaves, stems and roots of
monocotyledons, mostly grasses, or endophytic on plant tissues, lichens, and marine algae, occasionally infecting
humans, or isolated from soil. Sexual morph:
Pseudostromata visible as raised, linear, blackened areas on
the host surface, with neck and upper surface visible through
slits in the host tissue, mostly gregarious, fusiform, ellipsoid to
irregular, black, cells between ascomata and darkened layer
above usually thick, composed of brown cells of textura
Fig. 19 Ciferriascosea fluctamurum (holotype). a Ascostromata on
substrate. b Cross section. c Peridium. d Paraphyses. e–g Asci. h J+
apical ring of ascus mounted in Melzer’s reagent. i–m Ascospores. n
Upper surface of culture. o Lower surface of culture. Scale bars: a=
1 mm, b=50 μm, c–h =10 μm, i–m=5 μm
angularis. Ascomata solitary or usually gregarious in linear
groups, immersed under pseudostromata, globose to
subglobose, membranous, papillate, ostiole with periphyses.
Peridium composed of several layers, comprising smallish,
brown, reddish brown to hyaline cells of textura angularis.
Hamathecium composed of dense, hypha-like, long, broad,
septate, branched, paraphyses. Asci 8-spored, unitunicate,
broad cylindrical to clavate or subglobose, without an apical
apparatus. Ascospores overlapping 1–3-seriate to irregularly
arranged, hyaline, apiosporous, with a large, straight or
curved, upper cell and smaller lower cell, usually surrounded
by a gelatinous sheath. Asexual morph: Coelomycetous or
Hyphomycetous, Arthrinium spp. Conidiomata sporodochial,
solitary to gregarious, immersed, erumpent from host tissue
when mature, irregular, black, carbonaceous, coriaceous.
Conidiomata basal stroma composed of several layers of dark
brown to hyaline cells of textura angularis, with thick side
wall, thin at upper and lower walls. Setae absent, or occasionally present, intermingled among conidiophores.
Conidiophore mother cells ampulliform, verrucose wall, producing a single single hyphoid, cylindrical, 1–2-septate, verrucose, flexuous conidiophore. Conidiogenous cells basauxic,
cylindrical, with/without verrucose wall. Conidia globose to
subglobose, dark brown, smooth-walled or with minute wall
ornamentations, with a truncate basal scar.
Type: Apiospora Sacc., Atti Soc. Veneto-Trent. Sci. Nat.,
Padova, Sér. 4: 85 (1875),
Notes: Apiosporaceae was introduced by Hyde et al.
(1998), and is typified by Apiospora. This monophyletic family is characterized by apiospores, and a basauxic, Arthriniumlike conidiogenesis (Samuels et al. 1981; Hyde et al. 1998;
Bahl 2006). Basauxic conidiogenesis is not very common, but
is a unique type of ‘conidial development’ (Bhat 2010), seen
only in some conidial fungi. Examples include genera such as
Arthrinium (Apiosporaceae, Xylariales), Dictyoarthrinium
(Pezizomycotina incertae sedis), Cordella (Apiosporaceae,
Xylariales), Pteroconium (Apiosporaceae, Xylariales) and
Spegazzinia (Pezizomycotina incertae sedis) (Ellis 1971,
1976; Kirk et al. 2008; Wijayawardene et al. 2012).
However, these are not phylogenetically closely related (Liu
et al. 2015), as molecular data only exists for Arthrinium;
sequence data atributed to Spegazzinia in GenBank are probably wrong. In basauxic conidiogenesis, each conidiophore
consists of a short ‘conidiophore-mother-cell’ (i.e., basal cell,
in most cases ampulliform in shape) and an extensible filament arising from within it, which is conidiogenous. In this
‘conidiogenous filament’, the first conidium is always
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terminal and holoblastic. Sometimes, this terminal cell also
remains sterile (Bhat 2010). Later formed conidia are all of
the blastic-type, but develop laterally and basipetally (onebelow the other). Apiospora was previously placed in the
family Amphisphaeriaceae (Müller and von Arx 1962). Barr
(1976) transferred Apiospora to Hyponectriaceae and later
moved the genus to Lasiosphaeriaceae (Barr 1990; Barr and
Cannon 1994; Hawksworth et al. 1995). Hyde et al. (1998) reexamined the type species of Apiospora (A. montagnei Sacc.),
described a new taxon A. sinensis and introduced the family
Apiosporaceae. Phylogenetic analyses have shown that
Apiospora is a distinct group accommodated in the order
Xylariales (Huhndorf et al. 2004; Zhang et al. 2006;
Jaklitsch and Voglmayr 2012; Dai et al. 2014; Sharma et al.
2014). In Crous and Groenewald (2013), Arthrinium is morphologically and phylogenetically circumscribed, and the sexual genus Apiospora is treated as a synonym, on the basis that
Arthrinium is the older and more commonly encountered
name, and is more frequently used in literature.
Arthrinium subglobosa D.Q. Dai & K.D. Hyde, sp. nov.
Index Fungorum number: IF551172; Facesoffungi
number: FoF 00630; Fig. 20
Etymology: In reference to the subglobose asci.
Holotype: MFLU 15–0384
Saprobic on dead bamboo culms. Sexual morph:
Pseudostromata visible as raised, blackened areas on the host
surface, with parallel spots and upper black surface visible
through slits in the host tissue, 1–3 mm long, 300–450μm
wide, 150–300μm high, solitary to gregarious, in linear rows,
irregular, black, cells between ascomata and darkened layer
above usually 50–200μm thick, composed of brown to pale
brown, large, thin-walled cells of textura angularis mixed
with the host tissues above. Ascomata 200–350μm diam.,
150–200μm high, in groups of 3–5, immersed, subglobose
with a flattened base, dark brown, coriaceous, papillate,
ostiolate at the centre. Peridium laterally 10–20μm thick,
composed of dark brown, thick-walled cells of textura
angularis. Hamathecium composed of 3–10μm diam., long,
filamentous, septate, unbranched, tapering paraphyses. Asci
75–150 × 27–36 μm (x = 105 × 35.6 μm, n = 20), 8-spored,
unitunicate, broadly cylindrical, clavate to subglobose, pedicel
indistinct, apically rounded, with a shallow apical chamber.
Ascospores 24–28×8.5–12.5μm (x =25.8×9.5μm, n=20),
overlapping 1–3-seriate, hyaline, apiosporous, clavate to
sub-cylindrical, straight or curved, apical cell large, 17–22×
7–12.5μm, with smaller basal cell, 5–10×5–7μm, occasionally at an acute angle from the larger cell at the constricted
septum, smooth-walled, guttulate, surrounded by 5–8μm
thick gelatinous sheath. Asexual morph: Undetermined.
Culture characters: Ascospores germinating on PDA within 24 h and germ tubes produced from the lower cell. Colonies
fast growing on PDA, reaching 6 mm in 2 weeks at 28 °C,
circular, with an irregular edge, pale brown from above,
Fig. 20 Arthrinium subglobosa (holotype). a Appearance of
pseudostromata in linear rows. b, d Section through ascoma also
showing pseudostroma. c Asci. e Peridium of ascoma. f Paraphyses. g,
h Ascospores. i Germinating ascospore. j, k Culture on PDA, k from
below. Scale bars: a=1 mm, b=50 μm, c–e=10 μm, f–i=5 μm
brown to dark brown in the centre from below. Mycelium
superficial to immersed in the media, with branched, septate,
smooth hyphae.
Material examined: THAILAND, Chiang Rai, Mae Sae
Village, on dead culm of bamboo, 14 May 2011, Dong-Qin
Dai DDQ00034 (MFLU 15–0384, holotype), living culture,
MFLUCC 11–0397, KIB s44, GZAAC.
Notes: Arthrinium subglobosa is characterized by linear
pseudostromata and broadly cylindrical, clavate to subglobose
asci and ascospores which are occasionally sharply curved.
Our new strain can be compared with Apiospora bambusae
(Turconi) Sivan. and A. setosa Samuels et al. in having almost
similar sized asci and ascospores. However, A. subglobosa is
phylogenetically separated from A. bambusae and A. setosa
based on combined analysis of multigene sequence data
(Fig. 1). Other bambusicolous species of Apiospora do not
have subglobose asci and the size of asci is less than 27μm
wide (Theissen and Sydow 1915; Hyde et al. 1998; Bahl
2006). In Arthrinium subglobosa asci are more than 27μm
wide.
Arthrinium hyphopodii D.Q. Dai, D.J. Bhat & K.D. Hyde,
sp. nov.
Index Fungorum number: IF551173; Facesoffungi
number: FoF 00631; Fig. 21
Etymology: In reference to the conidia germ tubes producing hyphopodia
Holotype: MFLU 15–0383
Saprobic on dead bamboo culms. Sexual morph:
Undetermined. Asexual morph: Coelomycetous.
Pseudostromata solitary to gregarious, visible as raised,
pustulate, blackened areas on the host surface, somewhat linear, with black conidial masses produced through slits in the
host tissue. Conidiomata sporodochial, 300–500μm wide,
400–550μm long, 150–250μm high, solitary to gregarious,
immersed in host tissue, erumpent when mature, irregular,
black, coriaceous. Conidiomata wall 100–150μm thick, lower
wall thin, around 10–20μm wide, composed of one layer of
dark brown to brown to hyaline cells of textura angularis.
Conidiophore mother cells ampulliform, verrucose, producing
a single conidiophore. Conidiophores 5–15×2–3.5μm (x =
9.5×2.6μm, n=20), hyphoid, cylindrical, 1–2-septate, strait
to flexuous, verrucose. Conidiogenous cells 4–6×2–3.5μm
(x =5.5×2.6μm, n=20), basauxic, cylindrical, discrete, with
verrucose wall. Conidia 5–10×4–8μm (x =6.5×5.6μm, n=
20), globose to subglobose, dark brown, smooth-walled, with
a truncate basal scar, occasionally with a longitudinal, hyaline,
thin, germ-slit.
Fungal Diversity
Fungal Diversity
Fig. 21 Arthrinium hyphopodii (holotype). a Appearance of darkened
pseudostromata on bamboo host. b–d Pseudostromata on bamboo host
with linear cracking. e–g Sections of conidiomata. h Conidioma wall. i–l
Conidiophores and conidiogenous cells. m Dark brown conidia. n, o
Germinating conidia with hyphopodia. p, q Cultures on PDA. Scale
bars: a=5 cm, b=1 mm, c, d=500 μm, e–h=100 μm, l–p=5 μm, q, r=
25 mm
Culture characters: Conidia germinating on PDA within
24 h with germ tubes arising from the germ-slit, often
terminating in pale to dark brown, branched, lobed, irregular,
thick-walled hyphopodia. Colonies growing fast on PDA,
Fungal Diversity
reaching 90 mm in 2 weeks at 26 °C, cottony, circular, edge
irregular, white from above, brown to dark brown in centre
from below. Mycelium superficial to immersed in/on media,
with branched, septate, smooth, hyphae.
Materials examined: CHINA, Kunming, Kunming
Institute of Botany, Chinese Academy of Sciences, on dead
culm of Bambusa tuldoides Munro (Poaceae), 7 July 2014,
Dong-Qin Dai DDQ00280 (MFLU 15–0383, holotype); living culture, MFLUCC 15–003, CBS; Ibid. (isotype in KUN,
HKAS 83868).
Notes: Arthrinium was previously known as the asexual
morph of Apiospora and has more than 50 species listed in
Index Fungorum (2015). In Arthrinium conidiogenesis is
basauxic (Ellis 1971; Bhat 2010). This new taxon produces
brown hyphopodia at the apex of the germ tubes. Such a
character has not been observed in other species of
Arthrinium. Based on the blast search using the ITS sequence
in NCBI’s GenBank, the closest hits are species of
A. pterospermum with poor coverage because of the sequence
length. A similar search using the LSU sequence showed as
closest hits A. ovatum (KF144950; Identities = 867/885
(98 %), Gaps= 3/885 (0 %)), A. phragmites (KF144956;
Identities = 865/885 (98 %), Gaps = 3/885 (0 %)) and
A. rasikravindrii (KF144961; Identities=866/887 (98 %),
Gaps=5/887 (0 %)).
Cainiaceae J.C. Krug, Sydowia 30 (1–6): 123 (1978)
[1977]
Facesoffungi number: FoF 00687
Saprobic on dead grasses and other monocotyledons,
appearing as shiny black dots, slightly effuse from the substrate. Sexual morph: Pseudostromata poorly developed or
lacking, or sometimes clypeate, scattered, superficial, domeshaped or slightly effuse, smooth, dark brown to black.
Ascomata immersed, solitary or aggregated, globose,
subglobose or lenticular, coriaceous, brown, with papillate
ostiole. Ostiolar papilla short, internally lined by small, hyaline periphyses. Peridium one or two-layered, if two-layered,
outer layer comprising thick-walled, brown cells of textura
angularis and inner layer comprising thin-walled hyaline cells
of textura angularis. Hamathecium comprising abundant, filamentous, septate, paraphyses, slightly constricted at the septa
and longer than asci. Asci 8-spored, unitunicate, cylindrical,
broadly cylindrical to clavate, short pedicellate, with a complex, layered, J+, cylindrical or dome-shaped apical ring.
Ascospores uniseriate, overlapping uniseriate to biseriate, hyaline when young and very dark brown at maturity, ellipsoidal
or sphaerical, unicellular to 1-septate, slightly constricted at
the septum, wall ornamented with reticulations or longitudinal
striations, longitudinal germ slits or germ pores, surrounded
by a hyaline gelatinous sheath. Asexual morph:
Coelomycetous. Conidiomata pycnidial, scattered, immersed,
globose to subglobose, black. Conidiophores hyaline, denticulate, sympodially proliferating. Conidiogenous cells with 1–
3 phialides, filiform, branched or simple, septate, hyaline.
Conidia elongate fusiform, falcate to lunate, unicellular or
septate, hyaline, with pointed ends.
Type: Cainia Arx & E. Müll., Acta bot. neerl. 4 (1): 111
(1955)
Notes: The family Cainiaceae was introduced to accommodate species with a unique ascal apical ring, comprising a
series of rings and longitudinal germ slits in the ascospores
(Krug 1978). Almost all members in this family are from
monocotyledons and mostly grasses. In addition to Cainia,
Atrotorquata, Arecophila, Ceriophora, Reticulosphaeria and
Ommatomyces have been placed in this family at various
times (Kohlmeyer and Kohlmeyer 1993; Hyde et al. 1996).
Seynesia has also been included based on phylogenetic analysis (Maharachchikumbura et al. 2015). Sequence data are
available in GenBank for Arecophila, Cainia and Seynesia
and their placement in Cainiaceae is confirmed based on molecular analysis (Smith et al. 2003; Maharachchikumbura et al.
2015). A new Atrotorquata species was introduced as
A. spartii (Liu et al. 2015), however analysis of LSU gene
sequence data indicate that the phylogenetic placement of
A. spartii is not in Cainiaceae although the morphological
characters are similar. However combined LSU and ITS gene
analysis (Fig. 1) shows the phylogenetic placement of
Atrotorquata in Cainiaceae. Hence we include Atrotorquata
in Cainiaceae. Another new genus, Amphibambusa was introduced from bamboo in Liu et al. (2015) and LSU gene
analysis and morphology indicates the genus belongs to
Cainiaceae. The asexual morph of Cainiaceae observed
from a pure culture of Cainia desmazieresii has been
reported by Muller and Corbaz (1956) as Rhabdospora-like.
However, Krug (1978) was unable to obtain the asexual
morph from Cainia desmazieresii in culture (Kang et al.
1999b) and we have also been unsuccessful.
Type species: Cainia graminis (Niessl) Arx & E. Müll.,
Acta bot. neerl. 4 (1): 112 (1955)
Cainia Arx & E. Müll., Acta bot. neerl. 4 (1): 111 (1955)
Facesoffungi number: FoF 00688
Type species: Cainia graminis (Niessl) Arx & E. Müll.,
Acta bot. neerl. 4 (1): 112 (1955)
Basionym: Delitschia graminis Niessl, Notiz. Pyr.: 48
(1876)
Saprobic or pathogenic on Poaceae, forming leaf or stem
spots, appearing as black, shiny, raised, scattered dots on host
surface. Sexual morph: Pseudoclypeus indistinct, scattered,
rarely clustered, superficial, dome-shaped, effuse, dark brown
to black, each containing a single, or rarely several ascomata,
surface smooth. Ascomata immersed, scattered, solitary or
aggregated, globose to subglobose, coriaceous, with papillate
ostioles. Ostioles short, internally covered with hyaline, filamentous periphyses. Peridium comprising outer layer of
thick-walled brown cells and inner layer comprising thickwalled hyaline cells, both of textura angularis.
Fungal Diversity
Hamathecium comprising numerous, filamentous, unbranched, septate paraphyses, slightly constricted at the septa,
longer than asci. Asci 8-spored, unitunicate, cylindrical to clavate, short pedicellate, apex rounded, with a complex, layered,
J+, wedge-shaped to cylindrical, apical ring. Ascospores overlapping uniseriate to biseriate, dark brown, ellipsoidal, with
one, median septum, constricted at the septum, wall with longitudinal striations, surrounded by a gelatinous sheath.
Asexual morph: Coelomycetous. Conidiomata pycnidia,
scattered, immersed, globose to subglobose, black.
Conidiophores hyaline, denticulate. Conidiogenous cells with
1–3 phialides, filiform, simple or branched, septate, hyaline.
Conidia elongate-fusiform to filiform, falcate to lunate, with
holoblastic conidiogenesis, unicellular or septate, hyaline,
sometimes with pointed ends (description of asexual morph
form Krug 1977 and Kang et al. 1999a, b).
Notes: Members of Cainia mainly occur on grass hosts.
Cainia was introduced by von Arx and Müller (1955) based
on Delitschia graminis Niessl and re-named as Cainia
graminis. This genus is characterized by immersed ascomata,
cylindrical asci with a characteristic J+ apical apparatus with
series of rings, brown, two-celled, ascospores with longitudinal striations, and ascospores surrounded by a mucilaginous
sheath. Cainia presently comprises C. desmazieri C. Moreau
& E. Müll. and C. graminis (Krug 1978). Cainia deutziae E.
Müll. and C. cupula (Ellis) E. Müll. were excluded from the
genus and placed in Arecophila as Arecophila deutziae (E.
Müll.) You Z. Wang et al. (Wang et al. 2004) and
Discostroma as Discostroma cupulum (Ellis) M.E. Barr respectively (Barr 1993).
Cainia graminis (Niessl) Arx & E. Müll., Acta bot. neerl. 4
(1): 112 (1955)
Facesoffungi number: FoF 00689; Fig. 22.
Saprobic on leaves of Lolium sp., appearing as black dots
on leaf. Sexual morph: Pseudoclypeus scattered, rarely clustered, superficial, dome-shaped, effuse, dark brown to black,
each containing a single, or rarely several ascomata, with
smooth surface. Ascomata 330–480μm high, 350–515μm
diam., (x =400×450μm, n=20), scattered, immersed, globose
to subglobose, coriaceous, short papillate, brown, extending
slightly above the surface of the pseudoclypeus, with small
ostiole. Papilla short, internally covered with hyaline
periphyses. Peridium 23–35μm wide (x =29μm, n=20), comprising outer, thick-walled, dark brown cells of textura
angularis and inner, thick-walled, hyaline cells of textura
angularis. Hamathecium comprising 2.5–4μm wide (x =
3.5μm, n=20), filamentous, septate, hyaline, paraphyses.
Asci 170–185×20–30μm (x =180×23μm, n=20), 8-spored,
unitunicate, cylindrical to slightly clavate, short pedicellate,
narrowly rounded at the apex, apical apparatus distinct,
dome-shaped, horizontally divided into a series of 3–4 rings,
ring at the apex J-, basal rings J+. Ascospores 26–30×10–
11μm (x =27.5×10.5μm, n=20), biseriate, hyaline when
Fig. 22 Cainia graminis (reference specimen). a Appearance of
ascomata on host substrate. b Cross section of ascoma. c Peridium. d
Paraphyses. e Complex J+, apical ring in Melzer’s reagent. f–i Asci. j–
n Ascospores. o Germinating ascospore, germtubes coming from germslit. Note: apical pad of ascospore showing in k. Scale bars: a=500 μm,
b=100 μm, c, j–o=10 μm, d, e=5 μm, f–i=20 μm
young, becoming pale yellow-brown, and finally very dark
brown at maturity, ellipsoidal, broadly narrow and rounded
towards the ends, 1-septate, septum in centre, slightly constricted at the septa, with 8 longitudinal striations superficially
resembling ridges and an apical thickening or mucilaginous
pad of the wall at each end of the spore, surrounded by a
hyaline gelatinous sheath, germination via germ slit.
Asexual morph: Undetermined.
Culture characters: Colonies growing on MEA, reaching
1 cm within 7 days at 18 °C, fast growing, circular, with
smooth flat colonies, centre of colony white, woolly and colony margin with unbranched, hyaline, hyphae, tightly
adpressed to the media.
Materials examined: ITALY, Province of Forlì-Cesena
[FC], Santa Sofia, Corniolo, stem of Lolium temulentum L
(Poaceae), 25 September 2013, Erio Camporesi, IT 1462
(MFLU 15–0724, reference specimen designated here); living culture, MFLUCC 15–0540, MFLUCC 14–0987, ICMP;
Ibid. (MFLU 15–0724bis, MFLU 15–0724tris, duplicates of
reference specimen).
Notes: Morphological comparison and a Mega blast search
using the LSU sequence indicated the closest hit to our strain
is Cainia graminis (CBS 136.62, AF431949; Identities=880/
885 (99 %), Gaps=1/885 (0 %)). As well as morphological
characters of our strain were almost identical to Cainia
graminis which was illustrated by Krug (1978). However we
do not epitype this collection because the host and locality of
type material is not the same, and thus designate it as a reference specimen (sensu Ariyawansa et al. 2014).
Cainia anthoxanthis Senan., Camporesi & K.D. Hyde, sp.
nov.
Index fungorum number: IF551188; Facesoffungi number:
FoF 00690; Fig. 23.
Etymology: Named after the host genus on which the fungus occurs.
Holotype: MFLU 15–0539
Saprobic on leaves of Anthoxanthum odoratum L.,
appearing as black dots on leaf surface. Sexual morph:
Pseudoclypeus scattered, superficial, dome-shaped, thick, occasionally slightly effuse, black, each containing a single
ascomata, smooth. Ascomata 270–300μm high, 230–250μm
diam., (x =280×235μm, n=20), scattered, immersed, globose
to subglobose, coriaceous, brown, extending slightly above
the surface of the pseudoclypeus, ostiolate, papillate. Papilla
short, internally covered with hyaline, filamentous,
periphyses. Peridium 10–20μm wide (x =17μm, n=20) outer
Fungal Diversity
Fungal Diversity
Fig. 23 Cainia anthoxanthis (holotype) a Appearance of pseudoclypeus
on the host substrate. b Cross section of ascoma showing black
pseudoclypeus around the neck c Peridium. d Paraphyses. e–g Asci in
water. h–i Asci in Melzer’s reagent with J+ ring. j Apical J+ ring of ascus.
k–o Ascospores. p Ascosporic sheath in Indian ink. q Culture growing on
MEA from above. r Culture growing on MEA from below. Note: The
ascospore in p is mounted in Indian ink and after adding this chemical the
texture of the ascospore changed. Scale bars: a=500 μm, b=100 μm, c–
d=10 μm, e–p=50 μm
layers comprising of thick-walled, dark brown cells of textura
angularis and inner layers, of thin-walled, hyaline cells of
textura angularis. Hamathecium comprising 2.5–5.5μm wide
(x =3.6μm, n=20), filamentous, septate, hyaline, paraphyses.
Asci 170–205×30–40μm (x =180×35μm, n=20), sparse, 8spored, unitunicate, ovoid to clavate, slightly curved, with
indistinct short pedicel, narrowly rounded at the apex, apical
apparatus 6.5–8.5μm high, 6–9.5μm wide (x =8×7.5μm, n=
10), J+, distinct, dome-shaped, divide in two parts by vertical
furrows and horizontally divided into a series of 3–5 rings,
ring at the apex of the apparatus J-, basal rings J+. Ascospores
25–35×10–15μm (x =30×12μm, n=20) biseriate, hyaline
when young, becoming pale yellowish brown and finally very
dark brown at maturity, ellipsoidal, broadly narrow and
rounded at the ends, 1-septate, septum median, slightly constricted at the septum, with 8 longitudinal striations, superficially resembling ridges and an apical thickening of the wall at
each end of the spore, surrounded by a hyaline, gelatinous
sheath. Asexual morph: Undetermined.
Culture characters: Cultures on MEA slow growing,
reaching 1 cm diam. within 10 days at 18 °C, colonies circular,
flat, with diffuse margins. Centre of the colony white, woolly,
with clusters of mycelia, margin of the colony with unbranched, hyaline, mycelia.
Materials examined: ITALY, Province of Forlì-Cesena
[FC], Premilcuore, Valbura, on leaves of Anthoxanthum
odoratum L. (Poaceae), 2 February 2013, Erio Camporesi,
IT 1040 (MFLU15–0723, holotype); ex-type living culture,
Fungal Diversity
MFLUCC 15–0539, ICMP. Ibid. (MFLU 15–0575, isotype);
Ibid. MFLU15–0723bis, MFLU 15–0575bis, MFLU15–
0723tris, paratypes).
Notes: Based on a megablast search of the GenBank nucleotide database, the closest hit using the LSU sequence for our
collection (MFLU 15–0723) is Cainia graminis (CBS 136.62,
AF452033, Identities=900/930 (98 %), Gaps=0/930 (0 %)).
Morphologically Cainia anthoxanthis differs from Cainia
graminis in having small ascomata, asci with a biseriate ascospore arrangement, a shorter ascal apical ring with a rounded
apex and ascospores with a large sheath. The ascospores of
Cainia anthoxanthis are more oval at maturity and apical pad
is not obvious. However Cainia graminis having more elongated, fusiform ascospores with prominent apical pad. Based
on both morphology and molecular phylogeny, Cainia
anthoxanthis is identified as a new species of Cainia.
Coniocessiaceae Asgari & Zare, Mycol. Progr. 10 (2): 195
(2011)
Facesoffungi number: FoF 00671
Saprobic on grasses, soil and dung. Sexual morph:
Ascomata small, less than 200 μm diam., superficial,
subglobose to pyriform, glabrous or pilose, ostiolate, commonly with hypha-like ostiolar projections. Peridium thin,
membranaceous, translucent or sometimes opaque, outer layer
with cells of textura intricata. Hamathecium with abundant or
few paraphyses. Asci 4-spored, unitunicate, cylindrical to
subcylindrical, J-, without apical ring structures. Ascospores
dark brown to black, 1-celled, ellipsoidal or fusiform, smoothwalled, with a distinct germ-slit extending over the entire
length of spores. Asexual morph: Hyphomycetous,
nodulisporium-like. Conidiophores micronematous to
macronematous, simple or branched, smooth-walled or verrucose; hyaline. Conidiogenous cells integrated, terminal, discrete, elongating sympodially, with persistent conspicuous
denticles, hyaline. Conidia globose, subglobose to pyriform,
smooth-walled or verruculose, with a rounded apex, and hyaline, attenuated and truncated base and a more or less distinct
projection at the point of attachment to the conidiogenous
cells (Asgari and Zare 2011).
Notes: The family Coniocessiaceae was introduced by
Asgari and Zare (2011) to accommodate the genus
Coniocessia with a nodulisporium-like asexual morph.
García et al. (2006) had introduced the genus and placed it
in Xylariales genera, incertae sedis. The family is unique in
the order Xylariales in having a hyaline asexual morph with
polyblastic conidiogenesis (nodulisporium-like), contrasting
with the generally pigmented Nodulisporium asexual morphs
in Xylariales (Asgari and Zare 2011). Members of
Coniocessiaceae can also be distinguished from Xylariaceae
in possessing nonstromatic ascomata, and asci with nonamyloid apical structures (Asgari and Zare 2011). Asgari
and Zare (2011) introduced four new species in Coniocessia
and provided molecular data indicating this was a distinct
lineage of Xylariales. Thus they introduced the new family.
In this study (Fig. 1) we confirm its distinctness as a family
and place it in the order Xylariales.
Type: Coniocessia Dania García et al., in García et al.,
Mycol. Res. 110 (11): 1284 (2006)
Type species: Coniocessia nodulisporioides (D. Hawksw.)
Dania García et al., in García et al., Mycol. Res. 110 (11):
1285 (2006); Fig. 24.
≡ Coniochaeta nodulisporioides D. Hawksw., Norw. Jl
Bot. 25 (1): 15 (1978)
Diatrypaceae Nitschke [as ‘Diatrypeae’], Verh. naturh. Ver.
preuss. Rheinl. 26: 73 (1869)
Facesoffungi number: FoF 00679
Saprobic or pathogenic mostly on trees. Sexual morph:
Stromata eustromatic or pseudostromatic, well-developed,
immersed to erumpent or rarely superficial, mostly black or
dark brown, with somewhat carbonaceous outer layer, inner
layer pale, loosely packed, parenchymatous. Ascomata perithecial, immersed in stromatic tissues, globose to subglobose,
coriaceous, mostly brown, with long, ostiolar necks. Ostioles
sulcate, inner layer covered with hyaline, periphyses.
Peridium with two layers, outer layer comprising brown cells
of textura angularis and inner layer comprising hyaline cells
of textura angularis. Hamathecium comprising long, wide,
thin-walled, branched, septate, paraphyses, arising from base
of perithecia and slightly constricted at septa. Asci 8-spored or
polysporous, unitunicate, cylindrical, with a very long pedicel
with a more or less truncate apex, with J-, or J+ apical apparatus. Ascospores hyaline to light brown, crowded, allantoid.
Asexual morph: Coelomycetous, libertella-like.
Conidiomata on host, acervular, subcortical, astromatic,
erumpent, yellow to red, with branched conidiophores, in culture pycnidial, superficial, solitary or aggregated, subconical,
globose to subglobose, yellow, dark brown to black, with
thick peridium, comprising brown, thick-walled cells of
textura angularis with branched conidiophores, arising from
pseudoparenchymatous cells or interwoven hyphae.
Conidiogenous cells in dense palisades, cylindrical, straight
or curved, apically distorted or bearing annellations. Conidia
filiform, curved, or rarely straight, with flattened base and
blunt apex, hyaline.
Type: Diatrype Fr., Summa veg. Scand., Section Post.
(Stockholm) 384 (1849)
Type species: Diatrype disciformis (Hoffm.) Fr., Summa
veg. Scand., Section Post. (Stockholm): 385 (1849)
Notes: The family Diatrypaceae was introduced by
Nitschke (1869) and is a well-studied family comprising 15
genera (Maharachchikumbura et al. 2015). The type genus is
Diatrype, and other genera are Anthostoma, Cryptosphaeria,
Cryptovalsa, Diatrypella, Diatrypasimilis, Echinomyces,
Eutypa, Eutypella, Leptoperidia, Monosporascus,
Pedumispora, Peroneutypa, Phaeoisaria, and Quaternaria
(Maharachchikumbura et al. 2015). Sequence data is lacking
Fungal Diversity
Fungal Diversity
Coniocessia nodulisporioides (illustration based on Asgari and
Zare (2011) a Masses of ascospores on host surface. b Ascomata. c
Hypha-like ostiolar projections. d Paraphyses. e Immature ascus. f
Mature ascus. g, h Ascospores. i Conidiophores and conidiogenous
cells. j Conidia. Scale bars: a=100 μm, b=50 μm, c–j=10 μm
Fig. 24
for Echinomyces, Leptoperidia, Peroneutypa and
Quaternaria. Recent papers with sequence data providing
backbone trees are Carmaran et al. (2006) and Trouillas
et al. (2010) as well as Liu et al. (2015) in which a new species
of Diatrype from palms was introduced. In the present account
we provide a description and reference specimen (sensu
Ariyawansa et al. 2014) for the generic type, Diatrype
disciformis (Hoffm.) Fr. and a collection of Eutypa flavovirens
(Pers.) Tul. & C. Tul.
Diatrype Fr., Summa veg. Scand., Section Post.
(Stockholm) 384 (1849)
Facesoffungi number: FoF 00702
Saprobic on bark. Sexual morph: Stromata scattered or
aggregated on host substrate, sometimes spread over a large
area of the host, erumpent to superficial, orbicular, disc-like,
with flat or convex surface, sometimes spread over the host
evenly, arising through the cracks in bark epidermis or beneath
the epidermis, edges of cracks remaining as pointed, angular
parts, numerous ascomata immersed in one stroma, ostioles
opening to surface appearing as black spots, comprising an
outer, dark brown, small, tightly packed, thin parenchymatous
cell layer and inner, yellowish white, large, loosely packed,
parenchymatous cell layer. Ascomata perithecial, immersed in
stromatic tissues aggregated, globose to subglobose,
narrowing towards the apex and very narrow at the base of
ostiolar canal, pale brown, thin-walled, ostiolate. Ostiolar
canal immersed in only dark outer layer of stromata, short,
compressed, apex wider than base, periphysate, ostiolar opening covered with carbonaceous, black cells. Periphyses hyaline, filamentous, short, curved towards the cavity. Peridium
thin, comprising outer, brown, thick-walled cells of textura
angularis and inner, hyaline, thick-walled cells of textura
angularis. Hamathecium comprising long, wide, globose to
ovoid, thin-walled, branched, septate, paraphyses, slightly
constricted at septa, narrowing and tapering towards the blunt
apex. Asci 8-spored, unitunicate, long and narrow, with thinwalled pedicel, with cylindrical, thick-walled, swollen upper
portion, apex flat, with J-, cylindrical, conspicuous, apical
apparatus. Ascospores biseriate, hyaline, allantoid, unicellular,
with small, fat globules at each end, thin and smooth-walled.
Asexual morph: Coelomycetous, Libertella-like.
Conidiomata appearing as brownish yellow, watery, conidial
rounded mass on white, mycelial clumps. Pycnidia superficial, solitary or aggregated, subconical, globose to
subglobose, shiny, surface smooth, yellow, dark brown to
black. Peridium thick, comprising brown, thick-walled cells
of textura angularis. Conidiophores branched, arising from
pseudoparenchymatous cells or interwoven hyphae.
Conidiogenous cells in dense palisades, cylindrical, straight
or curved, apically distorted or bearing annellations. Conidia
filiform, curved or rarely straight, with flattened base and
blunt apex, hyaline.
Notes: Diatrype comprises mainly saprobes on decaying
wood. A few species are reported as pathogens forming cankers on forest trees. Species in this genus are quite resistant to
harsh conditions. The asexual morph of Diatrype is reported
as a libertella-like fungus (Adamčíková et al. 2011).
Diatrype disciformis (Hoffm.) Fr., Summa veg. Scand.,
Section Post. (Stockholm): 385 (1849)
Facesoffungi number: FoF 00691; Fig. 25.
Saprobic on wood with bark. Sexual morph: Stromata
1.5–2 mm wide, scattered on host, erumpent to superficial,
orbicular, disc-like, somewhat convex, arising through cracks
in bark epidermis, edges of cracks remaining as pointed, angular parts, with numerous perithecia immersed in a single
stroma, ostioles opening in the central region of the disc and
appearing as black spots, margin of the disk thick, black, composed of an outer, dark brown, small, tightly packed, thin
parenchymatous cell layer and inner, yellowish white, large,
loosely packed, parenchymatous cell layer. Ascomata 620–
760μm high, 265–360μm diam. (x =345×675μm, n=20),
perithecial, immersed in stromatic tissues, aggregated, globose to subglobose, narrowing towards the apex and very
narrow at the base of ostiolar canal, pale brown, thin-walled,
ostiolate. Ostiolar canal 60–105μm high, 85–140μm wide (x
=80×105μm, n=20), immersed in only dark outer layer of
stromata, short, compressed, apex wider than base,
periphysate, ostiolar opening covered with carbonaceous,
black cells. Periphyses hyaline, filamentous, short, curved towards the cavity. Peridium 20–30μm wide (x =25μm, n=20),
thin, comprising an outer layer of 5–7 layers of brown, thickwalled cells of textura angularis and a thin, inner strata of 7–
10 layers of hyaline, thick-walled cells of textura angularis.
Hamathecium comprising 70–75μm long, 1–2μm wide (x =
72×1.8μm, n=20), globose to ovoid, thin-walled, branched,
septate paraphyses, slightly constricted at septa, narrowing
and tapering towards the blunt apex. Asci 75–115μm long (x
=95μm, n=20), 8-spored, unitunicate, with very long, narrow,
thin-walled pedicel, with cylindrical, thick-walled, swollen
upper portion, 30–40×5–6μm (x =34×5.4μm, n=20), apex
flat, with J-, cylindrical, conspicuous, apical apparatus.
Ascospores 5–9×1.5–2μm (x =6.5×1.7μm, n=20), biseriate,
hyaline, allantoid, unicellular, thin-walled, with small, fat
globules at each end, smooth-walled. Asexual morph:
Undetermined.
Culture characters: Fast growing, reaching 4 cm within
7 days on MEA, when incubated at 18 °C, circular, with
smooth margin, fibrous, with sparse arial mycelium, white.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Santa Sofia Corniolino, on branch of Ostrya carpinifolia
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Fig. 25 Diatrype disciformis (reference specimen) a Stromata on
substrate. b Horizontal section of stroma. c Side view of stroma. d
Vertical cross section of stroma. e Cross section of ascomata. f
Peridium. g Ostiolar chamber filled with periphyses. h Asci
arrangement in hamathecium. i–k Asci. l Paraphyses. m Ascospores.
Scale bars: a=1 mm, b-d=500 μm, e, i–l=100 μm, g=50 μm, f, m=
10 μm
Scop. (Betulaceae), 14 November 2013, Erio Camporesi, IT
1516 (MFLU 15–0722, reference specimen designated
here); living culture, MFLUCC 15–0538, ICMP. Ibid.
MFLU 15–0722bis, duplicate of reference specimen).
Notes: Diatrype disciformis is the generic type of Diatrype
and represents the family Diatrypaceae. During our field survey, we collected D. disciformis associated with a branch of
Ostrya carpinifolia from Italy. We obtained herbarium
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specimens of D. disciformis from NY herbarium (NY305625,
NY305627 and NY305626) and compared the morphological
characters. The morphology of our fresh collection was similar to that of herbarium specimens. However, we could not
obtain the holotype specimen. Hence, we designate this as a
reference specimen (sensu Ariyawansa et al. 2014). The asexual morph of Diatrype disciformis is reported as Libertellalike, however we did not observe this in culture (Adamčíková
et al. 2011).
Eutypa Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 52
(1863)
Facesoffungi number: FoF 00703
Saprobic on decaying wood and pathogenic on living
plants causing dieback and cankers. Sexual morph:
Stromata scattered, erumpent to immersed, arising through
cracks in bark epidermis, several ascomata immersed in a
single stromata, ostioles opening to outer surface of stroma,
comprising an outer, black, small, tightly packed, thin parenchymatous cell layer and inner, whitish-yellow to dark yellow,
comprising of loosely packed, parenchymatous cells.
Ascomata perithecial, immersed in stromatic tissues, aggregated, globose to subglobose, narrowing towards the apex,
brown, thin-walled, ostiolate, papillate. Papilla short,
erumpent to immersed, conical, apex wider than base,
periphysate. Peridium comprising an outer layer of brown,
thick-walled cells of textura angularis and inner, hyaline,
thick-walled layer of cells of textura angularis.
Hamathecium comprising long, wide, septate, hyaline paraphyses, slightly constricted at septa, and narrowing and tapering
towards the apex. Asci 8-spored, unitunicate, long, narrow,
with thin-walled pedicel, with cylindrical, thick-walled, swollen upper portion, apex flat, with J-, cylindrical, conspicuous
apical apparatus. Ascospores biseriate or crowded, hyaline,
allantoid, unicellular, thin-walled, with small, fat globules at
the ends, smooth-walled. Asexual morph: Coelomycetous.
Conidiomata appearing as brownish-yellow, watery, bubblelike, rounded, conidial masses. Pycnidia superficial, solitary
or aggregated, subconical, globose to subglobose, shiny, with
smooth surface, yellow, dark brown to black. Peridium thick,
comprising brown, thick-walled, cells of textura angularis.
Conidiophores branched, arising from pseudoparenchymatous cells or interwoven hyphae. Conidiogenous cells cylindrical, in dense palisades, straight or curved, apically distorted
or bearing annellations. Conidia hyaline, filiform, curved or
rarely straight, with flattened base and blunt apex.
Notes: Eutypa comprises pathogens and saprobes and is
typified by Eutypa lata (Pers.) Tul. & C. Tul. Eutypa lata is
a serious pathogen on grapes forming cankers (Rolshausen
et al. 2004). Eutypa species are also very common on
decaying wood. However, the relationship between saprobic
and pathogenic species is not clarified.
Eutypa flavovirens (Pers.) Tul. & C. Tul., Select. fung.
carpol. (Paris) 2: 57 (1863)
Facesoffungi number: FoF 00692; Figs. 26 and 27.
Saprobic on decaying wood with bark. Sexual morph:
Stromata 1 –1.5 mm wide, scattered on host, erumpent, arising through the cracks in bark epidermis, cracks angular, with
numerous ascomata immersed in a single stromata, ostioles
opening to outer surface, appearing as black spots, comprising
an outer, dark brown to black, small, tightly packed, thin parenchymatous cell layer and inner, dark yellow, large, comprising of loosely packed, parenchymatous cells. Ascomata 295–
430μm high, 120–210μm diam., (x =330×180μm, n=20),
perithecial, immersed in stromatic tissues, aggregated, globose to subglobose, narrowing towards the apex and very
narrow at the base of papilla, pale brown, thin-walled,
ostiolate, papillate. Papilla 70–90μm wide, 85–120μm high
(x =70×105μm, n=20), short, immersed in only dark outer
layer of stromata, conical, apex wider than base, periphysate.
Peridium 11–16μm wide (x =13μm, n=20), comprising an
outer, brown, thick-walled, strata of 6–8 layers of cells of
textura angularis and inner, hyaline, thick-walled, strata of
4–6 layers of cells of textura angularis. Hamathecium comprising 3.5–5μm wide (x =4.5μm, n=20), long, septate, paraphyses, slightly constricted at septa, and narrowing and tapering towards the apex. Asci 140–210×9.5–13μm (x =170×
10.5μm, n=20), 8-spored, unitunicate, with narrow, long,
thin-walled pedicel, with cylindrical, thick-walled, swollen
upper portion, apex flat, with J-, cylindrical, 3.5–8.5μm high,
(x =5.6μm, n=20), conspicuous apical apparatus. Ascospores
7–11×1.5–2.5μm (x =8.3×2.1μm, n=20), biseriate, hyaline,
allantoid, unicellular, thin-walled, with small, fat globules at
ends, smooth-walled. Asexual morph: Mycelial clumps erect,
white, floccose on culture. Conidiomata 0.7–1.3 mm diam.,
appearing as brownish yellow, watery, bubble-like, rounded,
conidial masses forming from mycelial clumps. Pycnidia superficial, solitary or aggregated, subconical, globose to
subglobose, shiny, with smooth surface, yellow, dark brown
to black. Peridium thick, comprising brown, thick-walled,
cells of textura angularis. Conidiophores 4.5–5.5μm high,
2–2.5μm wide (x =4.9×2.4μm, n=20) branched, arising from
pseudoparenchymatous cells or interwoven hyphae.
Conidiogenous cells 9.5–13.5μm high, 2–3μm wide (x =
11.5 × 2.4 μm, n = 20), cylindrical, in dense palisades,
straight or curved, apically distorted or bearing
annellations. Conidia 30–40 × 1–2 μm (x = 34 × 1.4 μm,
n=20), filiform, curved or rarely straight, with flattened
base and blunt apex, hyaline.
Culture characters: Fast growing reaching 4 cm within
7 days on MEA, when incubated at 25 °C, circular, flat, with
diffuse margin, white, and becoming yellowish-white, bright
yellow to dull yellow with age.
Material examined: THAILAND, Chiang Rai Province,
Muang District, near Bandu, Baan Khuakhae, at 31 M. (19°
59’ 52.05” N; 99° 49’ 25.15” E), on decaying twigs, 15
November 2012, K.D. Hyde, CHUNI006 (MFLU15–0741,
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reference specimen designated here); living culture,
MFLUCC 13–0625, ICMP.
Notes: Fungal taxa in Eutypa are saprobes and some cause
serious cankers of forest trees (Trouillas et al. 2011). Eutypa
flavovirens was collected from northern Thailand and the
Libertella-like asexual morph formed in culture. Eutypa
flavovirens is morphologically distinct since it has yellow
stromatic tissues. There is no LSU sequence data available
for Eutypa flavovirens in GenBank. A BLAST searche of
ITS sequence data of this Eutypa flavovirens strain indicated
similarity to our Eutypa flavovirens (D13M, AJ302429;
Identities=545/572 (95 %), Gaps=12/572 (2 %)). The morphological characters of sexual and asexual morph of Eutypa
flavovirens are similar to those of previous studies (Glawe and
Rogers 1982; Carmarán et al. 2006; Trouillas et al. 2011) in
both morphology and phylogeny. However the specimen collected in Thailand may not be identical to the species which
was previously collected in Europe.
Graphostromataceae M.E. Barr et al., Mycotaxon 48: 533
(1993)
Facesoffungi number: 00624
Saprobic on trunks, branches, and twigs of members belonging to several genera of deciduous trees. Sexual morph:
Stromata widely effuse, irregular in outline, with two layered
ectostroma and entostroma, ectostroma develops first, young
ectostroma surface buff, with matted mycelium of yellow hyphae, ectostroma develops beneath, cinnamon in colour, hard
and brittle, composed of dark hyphae intermixed with suberized cells of the periderm, pulvinate in the centre. Ascomata
monostichous, immersed in diatrypoid configuration in
entostroma, globose, but compressed, bottle–shaped, more
rarely pyriform or obpyriform, varying in size with, black,
opaque, carbonaceous, doliiform perithecial embedded necks.
Peridium comprising hyaline, multi-layered, thin, indistinct,
flattened cells, about 10 μm wide. Hamathecium comprising
few, sparse, elongate, tapering, thin-walled paraphyses. Asci
unitunicate, 8-spored, fusiform to narrowly clavate, rounded
to subtruncate at the apex, with J+ apical apparatus.
Ascospores 1-celled, hyaline, multiseriately arranged,
suballantoid, tapering at both ends, thin-walled. Asexual
morph: nodulisporium-like (Barr et al. 1993)
Type genus: Graphostroma Piroz., Can. J. Bot. 52 (10):
2131 (1974)
Graphostroma Piroz., Can. J. Bot. 52 (10): 2131 (1974)
Facesoffungi number: 00704
Saprobic on trunks, branches, and twigs of members of
several genera of deciduous trees. Sexual morph:
Ectostroma cinnamon in colour in young stromata, hard and
brittle, composed of dark hyphae with periderm cells, at maturity ectostromatic crust separates exposing the entostromatic
“disc”, with ostioles, dark cinnamon to chestnut brown,
pulvinate in the centre, with a thin, sterile, fimbriate margin.
Ascomata more or less monostichous, compressed globose,
Fig. 26 Diatrype disciformis (reference specimen). a, b Stromata on
substrate. c Horizontal section of stroma. d Vertical section of stroma. e
Ostiolar chamber with periphyses. f Cross section of ascomata. g
Paraphyses. h Peridium. i–m Asci. n Ascospores. o Culture from
above. p Culture from below. Scale bars: a, b=500 μm, c=1 mm, d=
200 μm, e=50 μm, f=100 μm, g–n=10 μm
bottle-shaped, more rarely pyriform, or obpyriform, varying
in size from specimen to specimen. Peridium hyaline, composed of indistinct, tangentially flattened cells. Hamathecium
comprising hyaline, very thin-walled, inconspicuous narrowly
ventricose paraphyses, 3-septate, tapering gradually to a pointed apex. Asci 8-spored, unitunicate, clavate, subsessile to
short-stalked apex rounded to subtruncate, with a minute J+
apical apparatus. Ascospores overlapping biseriate, hyaline,
suballantoid, tapered at both ends, inequilateral, thin-walled,
with longitudinal germ-slit. Asexual morph: nodulisporiumlike. Conidiogenous cells thin-walled, often roughened at the
base, apex swollen into a clavate or globose vesicle, dilute
straw coloured towards the base, hyaline above, bearing up
to 12 small, inconspicuous denticles, arising singly from the
apex or sides of more or less erect mycelial hyphae, or in
whorls of three to five from apices of erect hyphae, Conidia
pyriform to turbinate, rounded above, hyaline, 1-celled,
strongly attenuated towards a minute thickened and refractive
basal scar, containing a single, large guttule.
Type species: Graphostroma platystoma (Schwein.) Piroz.,
Can. J. Bot. 52 (10): 2131 (1974)
Basionym: Diatrype platystoma (Schwein.) Berk.,
Grevillea 4 (no. 31): 95 (1876)
Facesoffungi number: FoF 00711; Fig. 28.
Notes: Pirozynski (1974) observed a North American species resembling Diatrype stigma (Hoffm.) Fr. and placed it in a
new monotypic genus Graphostroma, as G. platystoma
(Schwein.) Piroz. Pirozynski (1974) observed an association
between Graphostroma and a nodulisporium-like sp. in herbarium specimens and in culture which was identical to most
of the conidial states of Hypoxylon. Therefore Pirozynski
(1974) placed Graphostroma platystoma in Xylariaceae.
Barr and Boise (1985) positioned G. platystoma as atypical
in Xylariaceae and in Barr et al. (1993) introduced a new
monotypic family for Graphostroma based on the combination of its nodulisporium-like conidial state and diatrypaceous
sexual morph with allantoid ascospores. In the present phylogenetic analysis Graphostroma platystoma clusters together
with the species belongs to Hypoxylaceae (Fig. 1), while in
Maharachchikumbura et al. (2015) Graphostromataceae was
placed between Xylariaceae and Diatrypaceae.
The current, preliminary phylogeny based on rDNA data
raises some doubt as to whether the status of the
Graphostromataceae, as a separate family, will be supported
by molecular data. The species G. platystoma is defined by
having a sexual morph similar to that of Diatrypaceae, while
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Fungal Diversity
Fig. 27 Asexual morph of Eutypa flavovirens (MFLU 15–0741) a, b Arial clumps formed by mycelium. c, d Conidiomata on MEA. e Cross section of
conidioma. f Peridium. g–k Conidia attached to conidiophores. l–o Conidia. Scale bars: a–d=1 mm, e=100 μm, f=50 μm, g–o=10 μm
showing the characteristic asexual morph of the hypoxyloid
Xylariaceae. In the concept of Barr, which heavily relied on
the spore morphology, the family was therefore regarded as
intermediate between Diatrypaceae and Xylariaceae. On the
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Fig. 28 Graphostroma
platystoma (redrawn from
Pirozynski 1974). a Ascomata in
cross section. b Ascus and
paraphyses from DAOM 127673.
c Ascus from DAOM 75729. d
Ascospores from DAOM 36086.
e Ascospores from DAOM
65174. f Conidia. g
Conidiophores and conidia from
DAOM 134022. h Conidiophores
and conidia from DAOM 36091.
Scale bars: b–f=20 μm
Fungal Diversity
other hand, it has become clear that phylogenetic relationships
within the Xylariaceae are corroborated by the type of asexual
morph, and the current study clearly revealed a rather distant
relationship of Graphostroma with Diatrypaceae.
Morphologically similar xylariaceous genera with bipartite
stromata such as Biscogniauxia, Camillea and Obolarina will
need to be studied in detail to verify this hypothesis, using a
multigene genealogy.
Hyponectriaceae Petr., Annls mycol. 21 (3/4): 305 (1923)
Saprobic on dead plant matter. Sexual morph: Appearing
as black, shiny spots on host surface or small black lines
arising from cracks in bark. Pseudostromata present or lacking, if present, superficial, forming clypeus over the ascomata.
Ascomata solitary or aggregated, immersed, erumpent or rarely somewhat superficial, globose to ovoid, upright or horizontal, brown to black. Papilla short, ostiolate with or without
periphyses. Peridium comprising two layers, outer layer
of brown, cells of textura angularis, inner layer of hyaline cells of textura angularis, in some genera cells are
of textura globosa. Hamathecium of sparse, septate, paraphyses, tapering towards the apex, often deliquescing
at maturity. Asci 8-spored, unitunicate, oblong, cylindrical or ellipsoidal, short pedicellate, with a J+ or J- apical apparatus. Ascospores overlapping biseriate, or fasciculate, hyaline, yellow to light brown, fusoid,
isthmoid, elongate filiform, obovoid, or oblong, asymmetric or symmetric, unicellular or septate, smoothwalled or verruculose, with or without a mucilaginous
sheath. Asexual morph: Undetermined.
Type genus: Hyponectria Sacc., Michelia 1 (no. 2): 250
(1878)
Notes: Hyponectriaceae was introduced by Petrak (1923)
to accommodate both Hyponectria and Anisostomula.
Seventeen genera were included into the family
Hyponectriaceae by Hawksworth et al. (1995). Most genera
were, however, excluded from the family by Hyde et al.
(1998), while Palmomyces and Charonectria were added
(Hyde et al. 1998; Rossman et al. 1999). In addition to
Hyponectria, this family comprises of Apiothyrium,
Arecomyces, Arwidssonia, Cesatiella, Chamaeascus,
Discosphaerina, Exarmidium, Frondicola, Micronectria,
Pellucida, Physalospora, Phragmitensis, Pseudomassaria,
Rachidicola and Xenothecium (Wang and Hyde 1999;
Sivanesan and Shivas 2002). This is a problematic family
and it is not clear if it belongs in Xylariales. The type species
Hyponectria buxi (DC.) Sacc. needs recollecting and sequencing as the sequence data in GenBank may be inaccurate
(Jaklitsch and Voglmayr 2012). The other genera in this family
should also be recollected and sequenced to establish affinities. In this paper we include 17 genera in Hyponectriaceae.
Type species: Hyponectria buxi (DC.) Sacc.
Hyponectria buxi (DC.) Sacc., Michelia 1 (no. 2): 250
(1878)
≡ Sphaeria buxi DC., in de Candolle & Lamarck, Fl. franç.,
Edn 3 (Paris) 6: 146 (1815) Fig. 29.
Saprobic on leaves of Buxus sp. Ascomata immersed, depressed globose, visible as orange to brown dots on the host
surface, coriaceous, ostiolate, solitary or mostly gregarious.
Ostiole aperiphysate. Peridium one strata of 2 layers of cells
textura angularis with thickened brown walls around ostiole.
Paraphyses not observed. Asci 8-spored, unitunicate,
cylindric-clavate to clavate, short pedicellate, with an indistinct, J- apical apparatus. Ascospores overlapping biseriate or
obliquely uniseriate, hyaline, ellipsoidal or oblong, straight or
inequilateral, 1-celled, contents minutely guttulate, lacking a
sheath.
Material examined: UK, Surrey, Mickleham, on leaves of
Buxus sp., 1927, E.W. Mason no. 365 (IMI 16895).
Iodosphaeriaceae O. Hilber, in Hilber & Hilber, The
Genus Lasiosphaeria and Allied Taxa (Kelheim): 7 (2002)
Facesoffungi number: FOF 00849; Fig. 30.
Saprobic on host surface. Sexual morph: Ascomata superficial, solitary, black, and easily removed from the substrate,
ascomata covered with dark brown, setae-like hairs, comprising agglutinated mycelial strands, with a stellate arrangement,
arising from cells at the perithecial surface. Ostiole pore-like,
periphysate. Peridium comprising outer angular, pigmented,
brown cells, inner cells flattened, hyaline. Hamathecium comprising numerous, hypha-like, septate, flexuose, paraphyses,
slightly tapering towards the apex. Asci 8-spored, unitunicate,
narrowly clavate, short-pedicellate or apedicellate, apically
rounded, with a J+ subapical ring. Ascospores biseriate,
allantoid, unicellular, hyaline, smooth-walled, lacking sheaths
or appendages. Asexual morph: ceratosporium-like conidia
have been observed on the surface of perithecia, but may not
be related.
Notes: The family Iodosphaeriaceae was introduced by
Hilber and Hilber (2002) to accommodate the genus
Iodosphaeria. The genu s is unique in the order
Amphisphaeriales as it has superficial ascomata covered with
dark brown, setae-like hairs, comprising agglutinated mycelial
strands, with a stellate arrangement. Li et al. (2015b)
introduced a new species, Iodosphaeria tongrenensis Li
et al., bringing the total no. of species in the genus to
nine. Li et al. (2015b) provided molecular data that
showed this taxon clustered in Amphisphaeriaceae sensu
stricto and in this study (Fig. 1) we confirm its distinctness as a family.
Type: Iodosphaeria Samuels et al., Mycotaxon 28 (2): 486
(1987)
Type species: Iodosphaeria phyllophila (Mouton) Samuels
et al., Mycotaxon 28 (2): 486 (1987)
Material examined: NEW ZEALAND: Kaipara Harbour,
Mt Auckland, Atuanui State Forest; Cyathea dealbata,
November 1973, K.D. Hyde, GJS73-260 (PDD 32622,
holotype).
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Fig. 29 Hyponectria buxi
(redraw from plate in Wang and
Hyde (1999), IMI 16895) a
Ascomata on host substrate. b
Section of ascoma. c–e Asci. f–h
Ascospores. Scale bars: a–b=
40 μm, c–h=16 μm
Notes: When Samuels et al. (1987) introduced the genus
Iodosphaeria they deposited a specimen collected in New
Zealand in PDD and this was used in this study to illustrate
the family.
Lopadostomaceae Daranagama & K.D. Hyde, fam. nov.
Index fungorum number: IF551174; Facesoffungi number:
FoF 00071
Saprobic on dead wood. Sexual morph: Pseudostromata
(for Lopadostoma) immersed to erumpent, visible as darkened
areas, densely gregarious, scattered, compact. Stromata (for
Creosphaeria) erumpent to nearly superficial, with abrupt
margins, containing one to few perithecia, frequently coalescent in linear rows, dull black, waxy, roughened, flattened at
the top, inconspicuous perithecial mounds. Ascomata multiperithecial, single to multi-layered, arranged in a cluster, with
long ostiolar necks (for Lopadostoma), with ostiolar canal
lined with whitish material (for Creosphaeria). Ostioles umbilicate or at the same level as stromatal surface. Peridium
amorphous, comprising dark brown to black cells. Asci
(4–)8-spored, cylindrical, pedicellate, apically rounded, with
a J+ apical apparatus. Ascospores uniseriate or partly biseriate,
oblong, at first hyaline, turning light brown to nearly black,
narrowly ellipsoid, smooth-walled, with full length germ slit.
Asexual morph: Libertella-like. Conidiomata convoluted or
multi-loculate, with black pycnidial wall. Conidiophores dior trichotomously branched, stipes yellow, above hyaline,
smooth-walled, arranged in dense palisade. Conidiogenous
cells terminal, cylindrical, smooth-walled, bearing conidial
secession scars. Conidia hyaline, falcate, strongly curved to
semicircular, unicellular.
Type: Lopadostoma (Nitschke) Traverso, Fl. ital. crypt.,
Pars 1: Fungi. Pyrenomycetae. Xylariaceae, Valsaceae,
Ceratostomataceae 1 (2): 169 (1906)
Basionym: Anthostoma subgen. Lopadostoma Nitschke,
Pyrenomyc. Germ. 1: 121 (1867)
Notes: One of the hallmarks of most xylariaceous taxa are
the hyphomycetous asexual morphs. There are, however, certain exceptions where taxa produce coelomycetous asexual
morphs that have affinities with other families. In our phylogenetic analysis Lopadostoma was placed outside
Xylariaceae with Creosphaeria sassafras (Schwein.) Y.M.
Ju et al. and appears as a separate monophyletic group. The
same placement was also observed by Jaklitsch et al. (2014).
Lopadostoma and Creosphaeria are not very similar in
morphology and Ju et al. (1993) believe they should represent
two different genera. Jaklitsch et al. (2014) revised the genus
Fungal Diversity
Lopadostoma and also reported a Libertella asexual morph.
Creosphaeria was introduced by Theissen (1910) with having
soft, light coloured stromata. Petrini and Müller (1986) reported a Libertella asexual morph from culture. Ju et al. (1993)
observed the same type of conidia in the sexual morph.
Lopadostoma and Creosphaeria form separate monophyletic
groups, in a distinct, but previously unrecognized evolutionary line. Thus we introduce a new family, Lopadostomaceae,
in the order Xylariales, to accommodate Lopadostoma and
Creosphaeria. Creosphaeria may later require its own family
following more collections.
Lopadostoma (Nitschke) Traverso, Fl. ital. crypt., Pars 1:
F u n g i . P y r e n o m y c e t a e . X y l a r i a c e a e , Va l s a c e a e ,
Ceratostomataceae 1 (2): 169 (1906)
Facesoffungi number: FoF 00706
Saprobic on outer surface of wood and branches. Sexual
morph: Pseudostromata subglobose to conical, pustulate or
widely effuse, immersed, erumpent from bark, surrounded by
a narrow, black, carbonized encasement visible as “black
line”, containing one to many groups of perithecia,
ectostromatic disc visible, surrounded by blackened bark surface visible as a darkened area. Ectostromatic disc discrete to
fused, convex or pulvinate, flattened or raised, dark brown,
grey or black, smooth or tubercular. Ostiole inconspicuous
and umbilicate, rarely convex, shiny black, or distinctly
projecting and cylindrical. Ascomata clustered in valsoid
groups, single to double layers or rarely multi-layers,
subglobose or flask-shaped. Peridium amorphous, comprising
dark brown to black cells, with convergent ostiolar necks.
Hamathecium comprising numerous, long, apically free, rarely branched paraphyses. Asci 8-spored, unitunicate, cylindrical, pedicellate, with a globose to ellipsoid, J+ apical apparatus, containing a lower flat ring. Ascospores unicellular, oblong to narrowly ellipsoid, at first hyaline, becoming pale
brown and finally dark to blackish brown at maturity,
smooth-walled, germ slit straight through entire spore length,
with 2 large guttules when immature and a narrow hyaline
epispore. Asexual morph: Coelomycetous, Conidiophores
libertella-like, hyaline, rarely branched. Conidia unicellular,
hyaline, falcate, with acute upper end and narrowly truncate
lower end, curvature varying according to the species
(Jaklitsch et al. 2014).
Type species: Lopadostoma turgidum (Pers.) Traverso, Fl.
ital. crypt., Pars 1: Fungi. Pyrenomycetae. Xylariaceae,
Valsaceae, Ceratostomataceae 1 (2): 170 (1906)
Basionym: Sphaeria turgida Pers., Observ. mycol.
(Lipsiae) 1: 17 (1796)
Facesoffungi number: FoF00708; Fig. 31.
Notes: Lopadostoma turgidum was lectotypified by Lu and
Hyde (2000), which turned out to be invalid because the selected material was not from the original collection. Hence,
Jaklitsch et al. (2014) observed Persoon’s original material
(which is impossible to observe without actually going to L)
Fig. 30 Iodosphaeria phyllophila (holotype) a Herbarium packet. b
Herbarium material. c, d Ascomata on the host substrate. e Section of
ascoma. f Peridium. g Ceratosporium conidia seen from the surface of
perithecia. h–i J+ subapical ascal ring in Melzer’s reagent. j–m Asci (m
stained in Melzer’s reagent). n–q Ascospores. Scale bars: c=1 mm, d=
500 μm, e=50 μm, f=10 μm, g=20 μm, h–i=5 μm, j–m=10 μm, n–q=
5 μm
and epitypified Lopadostoma turgidum with a fresh
collection.
Material examined: AUSTRIA, Niederösterreich, Gaaden,
3 December 2011, H. Voglmayr (WU 32085, epitype).
Melogrammataceae G. Winter [as ‘Melogrameae’],
Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 797 (1886)
Facesoffungi number: FoF 00840
Saprobic on woody plants. Sexual morph: Stromata solitary, scattered or aggregated in lines, superficial, erumpent
from bark, pulvinate or discoid, soft textured, internally comprising reddish-brown, pseudoparenchymatous cells, apex papillate with black ostiole. Ascomata perithecial, sometimes
confluent, immersed, globose, medium, dark brown to black,
ostiole periphysate. Peridium comprising rows of cells, brown
externally, hyaline internally. Hamathecium of septate, hyaline, paraphyses. Asci 8-spored, unitunicate, clavate or fusoid,
straight, curved or sigmoid, short pedicellate, with shallow,
nonamyliod, apical ring. Ascospores hyaline or brown, filiform, straight or cylindrical or falcate, 0–3-septate, end cells
slightly entirely lighter or only at their tips, end narrowly
rounded to subacute, with a smooth narrow hyaline perispore,
often with one large guttule in each cell, wall smooth. Asexual
morph: Coelomycetous. Conidiophores long, cylindrical,
stiffly upright, septate, verticillate and whorled or not.
Conidiogenous cells holoblastic, proliferating sympodially,
hyaline, thin. Conidia elongate, falcate.
Type: Melogramma Fr., Summa veg. Scand., Section Post.
(Stockholm): 386 (1849)
Type species: Melogramma campylosporum Fr., Summa
veg. Scand., Section Post. (Stockholm): 386 (1849)
Notes: Melogramma was introduced by Fries (1849), with
Melogramma campylosporum as the type species. The family
Melogrammataceae was introduced by Winter (1884–1886)
to accommodate Botryosphaeria, Endiothia, Valsaria and
Melogramma. This family is characterized by pulvinate or
discoid stromata, with papillate black ostiolar dots, fusoid,
falcate, curved or sigmoid asci, and often strongly curved
ascospores (Jaklitsch and Voglmayr 2012).
Melogramma campylosporum Fr., Summa veg. Scand.,
Section Post. (Stockholm): 386 (1849)
Facesoffungi number: FoF 00841, Fig. 32
Basionym: Sphaeria melogramma (Bull.) Pers., Syn.meth.
fung. (Göttingen) 1:13 (1801).
Possible synonyms (see Index Fungorum 2015)
Saprobic on dead shoots of Carpinus betulus L. Sexual
morph: Stromata 545–694μm high, 1047–1668μm diam.,
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Fungal Diversity
solitary, scattered or aggregated in lines, superficial, erumpent
from bark, pulvinate or discoid, soft textured, reddish-brown,
internally comprising pseudoparenchymatous cells, apex papillate with black ostioles. Ascomata perithecial, (138–) 163–
287 (−297) μm high×(73–) 135–278 (−284) μm diam. (x =
225×206μm, n=10), sometimes confluent, immersed, globose, dark brown to black, ostiolate. Ostiole 54–72μm diam.,
with periphyses. Peridium 16–42μm wide, comprising two
layers, outer layer, composed of dark brown to dark cells of
textura angularis to prismatica, inner layer hyaline, with thin–
walled cells. Hamathecium of 3–4μm diam., septate, paraphyses. Asci (82–) 93–119 (−132)×(10–) 11–14 (−15) μm (x =
106×13μm, n=30), 8-spored, unitunicate, clavate or fusoid,
straight, curved or sigmoid, short pedicellate, with shallow, Japical ring. Ascospores (23.5–) 33–42 (−46)×(3.5–) 4–5 (−6)
(x =37.6×4.7μm, n=50), young ascospores hyaline, straight
and 0–1-septate, mature ascospores brown, falcate, often
strongly curved, 3-septate, end cells slightly entirely lighter
or only at their tips, ends narrowly rounded to subacute, with
a smooth, narrow, hyaline perispore; often with one large
guttule in each cell, smooth-walled.
Material examined: AUSTRIA, Styria, Graz, distr.
Mariatrost, on the north-facing hillside below the church,
clearing, 47°06’29”N, 15°29’32”E, MTB 8858/4, c. 440 m
alt.; 5 November 2006, C. Scheuer #5321 (S, F123341).
Pseudomassariaceae Senan. & K.D. Hyde, fam. nov.
Index fungorum number: IF551208; Facesoffungi number:
FoF 00843
Saprobic on recently dead twigs attached to the trees,
appearing as black dots, below small raised areas of bark,
opening through long fissures. Sexual morph: Ascomata
perithecial, scattered, solitary or aggregated, immersed, depressed globose to ellipsoid, coriaceous, black, ostiolate, papillate. Ostiolar papilla short, cylindrical, wide at the apex,
periphysate. Peridium comprising strongly compressed, narrow, light to medium brown, thin-walled cells of textura
angularis. Hamathecium comprising numerous, apically narrow, basally wide, hyaline paraphyses. Asci 8-spored,
unitunicate, clavate to fusoid, short pedicellate, apically
rounded, with J+, or J– apical ring. Ascospores biseriate or
partially uniseriate, hyaline, broadly ellipsoid, oblong or narrowly clavate, mostly apiosporous, with a rounded to
subconical, small, lower cell or equally uniseptate, straight
or curved, thick-walled, smooth-walled. Asexual morph:
Hyphomycetous, Setae erect, dark brown, straight to flexuous,
arising from superficial hyphae, branched at base,
subcylindrical, tapering to obtuse apex, 3–7-septate, basal cell
slightly swollen. Conidiophores subcylindrical to setiform,
with radially lobed basal cells, pale brown, smooth, arising
from superficial mycelium, straight to flexuous, 1–10-septate.
Conidiogenous cells terminal or lateral, polyblastic,
subcylindrical to somewhat clavate, pale brown, smooth, with
1–4 denticulate loci. Conidia lageniform, distal end free,
Fig. 31 Lopadostoma turgidum (epitype). a Herbarium specimen. b
Pseudostromata in face view. c Ectostromatic disc. d, e Transverse
pseudostroma sections. f, g Vertical pseudostroma sections. h Apical
ring bluing in Melzer’s reagent. i Peridium. j Paraphyses. k–o Asci.
p,r,s Ascospores showing germ slit. q Ascospore. Scale bars: b =
500 μm, c–e=200 μm, f–g=100 μm, h, j=10 μm, I=20 μm, k–o=
50 μm, p–s=5 μm
truncate, rostrate at proximal end, pale brown, with a
subhyaline transverse band at equatorial zone, smooth,
guttulate, aseptate.
Notes: Pseudomassaria was introduced by Jaczewski
(1894) to accommodate Sphaeria chondrospora. This genus
has been synonymised under Aplacodina, Apiospora and
Apiosporella (Barr 1964). Hyde et al. (1998) however,
assigned this genus to family Hyponectriaceae introducing a
new species as Pseudomassaria huwerae. Lumbsch and
Huhndorf (2010) also listed Pseudomassaria under
Hyponectriaceae as did Maharachchikumbura et al. (2015).
Combined gene analysis herein (Fig. 1), show the taxonomic
placement of Pseudomassaria different from
Hyponectriaceae. This result confirms the taxonomic placement suggested by Jaklitsch and Voglmayr (2012). The asexual morph of Pseudomassaria carolinensis M.E. Barr &
Hodges was reported as Beltraniella portoricensis (F.
Stevens) Piroz. & S.D. Patil) (Hodges and Barr 1971). All
species of Pseudomassaria species do not have similar morphological characters and phylogenetically they do not cluster
together. Leiosphaerella also shows molecular and morphological similarities with Pseudomassaria (Jaklitsch and
Voglmayr 2012). Single gene sequence analyses of ITS and
LSU regions in Jaklitsch and Voglmayr (2012) indicate a separate taxonomic grouping for the Pseudomassaria,
Leiosphaerella clade. Therefore in this study we introduce a
new family Pseudomassariaceae to accommodate
Leiosphaerella and Pseudomassaria.
Type: Pseudomassaria Jacz., Bull. Herb. Boissier 2: 663
(1894)
Type species: Pseudomassaria chondrospora (Ces.) Jacz.,
Bull. Herb. Boissier 2: 663 (1894)
≡ Sphaeria chondrospora Ces., Hedwigia 1: tab. 11, Fig. 2
(1855)
Pseudomassaria Jacz., Bull. Herb. Boissier 2: 663 (1894)
Facesoffungi number: FoF 00844
Saprobic on dead twigs attached to trees, appearing as
black dots, below small bumps on the bark. Sexual morph:
Ascomata perithecial, scattered, solitary or aggregated, immersed, depressed globose, coriaceous, black, ostiolate, papillate. Ostiolar papilla short, cylindrical, wide at the apex, even
with the bark surface, periphysate. Peridium comprising narrow, compressed, light brown, thin-walled, cells of textura
angularis at the base and thick-walled, light brown to hyaline,
isodiametric cells around the ostiole. Hamathecium comprising numerous, apically narrow, basally wide, hyaline
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paraphyses. Asci 8-spored, unitunicate, clavate to fusoid, short
pedicellate, apically rounded, with a J+ or J– apical ring.
Ascospores biseriate or partially uniseriate, hyaline, broadly
ellipsoid, oblong or narrowly clavate, apiosporous with a
rounded to subconical small lower cell, straight or curved,
thick-walled, smooth-walled. Asexual morph:
Hyphomycetous, Setae erect, dark brown, straight to flexuous,
arising from superficial hyphae, branched at base,
subcylindrical, tapering to obtuse apex, 3–7-septate, basal cell
slightly swollen. Conidiophores subcylindrical to setiform,
with radially lobed basal cells, pale brown, smooth, arising
from superficial mycelium, straight to flexuous, 1–10-septate.
Conidiogenous cells terminal or lateral, polyblastic,
subcylindrical to somewhat clavate, pale brown, smooth, with
1–4 denticulate loci. Conidia pale brown, lageniform to rhomboid, distal end free, proximal end rostrate, with a subhyaline
transverse band at equatorial zone, smooth-walled, guttulate,
aseptate.
Pseudomassaria chondrospora (Ces.) Jacz., Bull. Herb.
Boissier 2: 663 (1894).
Facesoffungi number: FoF 00845; Fig. 33.
Saprobic on dead twigs of Tilia cordata Mill.., appearing as
black spots, with long fissures on the host surface. Sexual
morph: Ascomata 610–625μm high, 765–800μm diam., (x
=620×775μm, n=20) solitary or aggregated, scattered, immersed, depressed globose to ellipsoid, brown, short papillate,
ostiolate. Ostiolar papillashort, cylindrical, wide at the apex,
even with the bark surface, inner layer of ostiolar canal covered with hyaline, periphyses. Peridium 25–45μm wide, (x =
45μm, n=20), comprising 5–7 layers of narrow, compressed, light brown, thin-walled cells of textura
angularis at the base and thick-walled, light brown to
hyaline, isodiametric ce lls around the ostiole.
Hamathecium comprising 10–17 μm wide (x = 12 μm,
n = 20), septate, branched paraphyses. Asci 200–250 ×
70–80μm (x =226×77μm, n=20), 8-spored, unitunicate,
broadly ellipsoidal, clavate to fusoid, short pedicellate,
apex rounded, with an indistinct, J- apical apparatus.
Ascospores 65–100 × 20–35 μm (x = 82× 26.5 μm, n =
20), biseriate or partly obliquely uniseriate, hyaline,
broadly ellipsoid, oblong or clavate, apiosporous, with
a rounded to subconical, small lower cell, straight or
rarely curved, thick-walled, smooth-walled, granulate.
Asexual morph: Undetermined.
Culture characters: Cultures growing on MEA reaching
1 cm. diam. at 18 °C, after 10 days. Slow growing, irregular
colony, margin cream yellow, central region gray and tightly
adpressed to the media, outer region woolly, serrate to smooth
margin.
Material examined: ITALY, Province of Forlì-Cesena,
Premilcuore, Fiumicello, on branch of Tilia cordata, 24
April 2013, Erio Camporesi, IT 1200 (MFLU 15–0729, reference specimen designated here); living culture, MFLUCC
Fig. 32 Melogramma campylosporum (S, F123341). a Herbarium
packet. b Host. c, d, f Stromata on wood surface. e Periphyses of
ascomata. g Peridium. h–l Asci. m Paraphyses. n–q Ascospores. Scale
bars: c, d, f=500 μm, e, g–m=30 μm, n–q=15 μm
15–0545, ICMP. Ibid. MFLU 15–0729bis, MFLU 15–
0729tris.
Notes: Pseudomassaria chondrospora is the generic type
of Pseudomassaria. However we could not locate the type
material of Pseudomassaria chondrospora and there is no
information of host and locality. However, Jaklitsch and
Vo g l m a y r ( 2 0 1 2 ) i l l u s t r a t e d a n d d e s c r i b e d t h e
Pseudomassaria chondrospora (WU 31321) collected from
twigs of Tilia platyphyllos and we compared morphological
characters of our collection with that illustration. As well as
Mega blast search results for LSU of our strain gives closest
match as Pseudomassaria chondrospora (PC1, JF440982;
Identity=1055/1097 (96 %), Gaps=20/1097 (1 %)). Hence
considering both morphology and phylogeny and lacking of
information about host and locality, we designate our collection as a reference specimen of P. chondrospora (sensu
Ariyawansa et al. 2014).
Pseudomassaria corni (Sowerby) Arx, Ber. schweiz. bot.
Ges. 62: 349 (1952)
Basionym: Sphaeria corni Sowerby, Col. fig. Engl. Fung.
Mushr. 3: pl. 370 (1803)
Facesoffungi number: FoF 00846; Fig. 34
Saprobic on dead twigs of Cornus sanguine L., appearing
as black, shiny, dots on host surface. Sexual morph:
Ascomata 195–240μm high, 280–315μm diam., (x =222×
299μm, n=20), solitary or aggregated in groups, scattered,
immersed, globose, brown, ostiolate. Ostiole short, wide, cylindrical. Peridium comprising two stratum, outer 5–6 layers
of dark brown, cells of textura globulosa, inner 1–2 layers of
narrow, compressed, hyaline to light brown, thin-walled, cells
of textura angularis. Hamathecium comprising very few, hyaline, septate paraphyses. Asci 55–80×12–15μm (x =67×
13μm, n=20), 8-spored, unitunicate, clavate to fusiform, short
pedicellate, apex rounded with indistinct, J- apical apparatus.
Ascospores 16.5–17.5×6–7.5μm (x =17×6.5μm, n=20),
biseriate to multiseriate, hyaline, oblong or clavate, ends
pointed, unicellular, thick-walled, smooth-walled. Asexual
morph: Undetermined.
Culture characters: Cultures growing on MEA at 18 °C,
after 14 days reaching 2 cm, with circular, yellowish white,
colonies, tightly adpressed to the media, woolly, loosely arranged, margin smooth.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Santa Sofia, Camposonaldo, on Cornus sanguine, 5
May 2013, Erio Camporesi, IT 1219 (MFLU 15–0728, reference specimen designated here), living culture, MFLUCC
15–0544 = ICMP. Ibid. MFLU 15–0728bis, MFLU 15–
0728tris, paratypes).
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Fig. 33 Pseudomassaria chondrospora (reference specimen). a, b Appearance of ascomata on host substrate. c Paraphyses. d Cross section of ascoma. e
Peridium. f–i Asci. j–p Ascospores. q Culture from upper surface. r Culture from lower surface. Scale bars: a, b=500 μm, d=100 μm, c, e–p=10 μm
Vialaeaceae P.F. Cannon, Mycol. Res. 99 (3): 368 (1995)
Facesoffungi number: FoF 00686
Biotrophic, parasitic or endophytic on dead plant matter,
often on dead twigs attached to trees. Sexual morph:
Pseudostroma (pseudoclypeus) absent or if present, immersed, ellipsoidal, as slightly raised black dots on substrate.
Ascomata perithecial, solitary or aggregated, immersed, globose to subglobose, coriaceous, ostiolate, with immersed,
Fungal Diversity
Fig. 34 Pseudomassaria corni (reference specimen) a, b Ascomata on substrate. c Cross section of ascoma. d Peridium. e–h Asci. i–m Ascospores.
Scale bars: a=500 μm, b=200 μm, c=20 μm, d–m=10 μm
periphysate, ostiolar canal, sometime converging and fusing
into the centre of the pseudostroma and opening through a
common ostiole, black to brown. Papilla long, sometimes
distal end curving towards the substrate or immersed ostiolar
canal Periphyses hyaline, filamentous. Peridium comprising
several layers of slightly flattened, strongly melanized, thickwalled cells of textura globulosa or textura angularis with
peridial pores between cells. Hamathecium comprising few,
filiform, rarely branched, septate, hyaline, paraphyses sometimes apically swollen and shorter than asci. Asci 8-spored,
unitunicate, cylindrical, apex rounded, short pedicellate, apex
obtuse or truncate, with a J+ sub-apical ring. Ascospores
biseriate, triseriate to fasciculate, sometimes weakly helically
coiled, 1–3-septate, hyaline, with two fusiform end cells, tapering gradually to acute ends and towards the central portion,
joined by a narrow, long isthmus. Asexual morph:
Ceolomycetous. Conidiomata pycnidia, superficial, solitary,
scattered, globose, with slimy, shining spore mass and basal
mycelium forming thick, black strands. Conidiophores erect,
branched, septate, hyaline. Conidiogenous cells phialidic, integrated, in small whorls, lageniform to cylindrical, hyaline.
Conidia oblong to ellipsoidal, one-celled, hyaline smoothwalled, with a truncate abscission scar.
Type: Vialaea Sacc., Bull. Soc. mycol. Fr. 12: 66 (1896)
Notes: Vialaeaceae was used by Schrantz (1960) as
“Vialaeacees” and formally established as Vialaeaceae by
Cannon (1995). This is a monotypic family comprising
Vialaea (Maharachchikumbura et al. 2015). Fungal members
of this family are endophytes or weak pathogens causing cankers on young twigs and on older branches of trees
(Senanayake et al. 2014).
Vialaea Sacc., Bull. Soc. mycol. Fr. 12: 66 (1896)
Fungal Diversity
Facesoffungi number: FoF 000011
Biotrophic, parasitic or endophytic in twigs of trees, eventually saprobic. Sexual morph: Pseudostromata lacking or if
present, immersed in host tissues, with brown to black tissues
around immersed neck. Ascomata perithecial, immersed, solitary or aggregated in circular groups with ostioles converging
in their centre, globose or subglobose, coriaceous, somewhat
flattened, black to brown. Ostiolar canal elongate, dark brown
to black, periphysate. Peridium composed of several layers of
slightly flattened, strongly melanized, thick-walled cells of
textura globulosa or textura angularis with peridial pores between cells. Hamathecium comprising poorly developed,
thin-walled, hyaline, septate, paraphyses, shorter than the asci.
Asci 8-spored, unitunicate, cylindrical, sometimes tapering towards the apex or base, short pedicellate or sessile, thin-walled
except at the apex, apex obtuse or truncate, subapical ring, J+,
subconical. Ascospores biseriate, triseriate to fasciculate,
sometimes weakly helically coiled, 1–3-septate, hyaline, with
two fusiform end cells, joined by a narrow, long isthmus.
Asexual morph: Coelomycetous, Conidiomata pycnidial, superficial, solitary, globose. Conidiophores erect, branched,
septate, hyaline. Conidiogenous cells enteroblastic,
annelledic, proliferating percurrently to produce small, hyaline, ellipsoidal conidia. Conidia oblong to fusiform, onecelled, hyaline and smooth-walled.
Vialaea mangiferae Senan. & K.D. Hyde, Sydowia 66 (1):
(2014); Fig. 35
Notes: This species was introduced in Senanayake et al.
(2014) and we illustrate both sexual and asexual morph here.
Material examined: THAILAND, Chiang Rai Province,
Muang District, near Bandu, Baan Khuakhae, at 31 M. 17,
(19° 59’ 52.05” N; 99° 49’ 25.15” E), on twigs of Mangifera
indica, 15 November 2012, K.D. Hyde, CHUNI001 (MFLU
13–0342, holotype.
Notes: Vialaea was introduced and is typified by Vialaea
insculpta (Fr.) Sacc. (Saccardo 1896) and placed in
Amphisphaeriaceae (Xylariales) based on the J+ apical ring
(Shoemaker et al. 2013). Cannon (1995) excluded Vialaea
from Amphisphaeriaceae and proposed a new family,
Vialaeaceae, for Vialaea species, based on the stromata,
ascomata and ascal characteristics, as well as nutritional strategy as a weak pathogen. Vialaea comprises of V. insculpta
(Fr.) Sacc., V. minutella Petr., and Vialaea mangiferae
Senanayake & KD. Hyde (Senanayake et al. 2014). Vialaea
bambusae Hara and V. ingae Rehm were excluded from the
genus by Cannon (1995). The above placements were based
on morphology and LSU gene sequences analysis has also
shown that Vialaeaceae is a distinct family in the order
Xylariales (Shoemaker et al. 2013; McTaggart et al.
2013; Senanayake et al. 2014; Maharachchikumbura
et al. 2015).
Xylariaceae Tul. & C. Tul. [as ‘Xylariei’], Select. fung.
carpol. (Paris) 2: 3 (1863)
Index fungorum number: IF81528, Facesoffungi number:
FoF00070
Saprobic, pathogenic, parasitic or endophytic in wood,
leaves and fruits, dung inhabitants, and associated with insect
vectors. Sexual morph: Stromata extremely variable in size,
shape and colour, erect or applanate or effused–pulvinate or
sometimes rudimentary, arising singly or aggregated, with 1 to
many ascomata, ostiolate, with or without extractable
stromatal pigments, bipartite or unipartite. Ascomata variable
in size and shape, globose-pyriform, embedded in the stroma,
single or multi-layered. Hamathecium comprised of hyphoid,
filamentous, septate paraphyses embedded in a gelatinous matrix. Asci 4–8-spored, unitunicate, cylindrical to clavate, pedicellate, apically rounded, with or without a J+ or J– apical
apparatus or with apical thickenings. Ascospores uniseriatebiseriate (except for the genus Phylacia), brown to black,
rarely hyaline, 1–2-celled, variously-shaped, mostly with a
germ slit. Perispore dehiscent or lacking smooth or with patterns. Asexual morph: Two major types of asexual morph
reported are nodulisporium-like and geniculosporium-like as
defined by Bitzer et al. (2008), featuring hyaline to light
brown, smooth, branched conidiophores bearing hyaline,
roughened or smooth, ellipsoidal conidia. Several other hyphomycetous and coelomycetous genera have been linked to
Xylariaceae (Crous et al. 2014).
Type: Xylaria Hill ex Schrank, Baier. Fl. (München) 1: 200
(1789).
Type species: Xylaria hypoxylon (L.) Grev., Fl. Edin.: 355
(1824)
Possible synonymy: See Index Fungorum (2015)
Notes: The family Xylariaceae was introduced by Tulasne
and Tulasne (1863) using the term “Xylariei”. The rank was
not certain as it did not address the family concept (Stadler
et al. 2014). Numerous studies have since contributed to the
knowledge of the family (i.e., Dennis 1957, 1961; Læssøe
et al. 1989; Miller 1961; Ju and Rogers 1996) and recent
studies have revised the family using ultra-structural data of
ascal and ascospore characters. Chemotaxonomic data have
also played an important role in the identification and characterization of the stromatic Xylariaceae (Whalley and Edwards
1995; Bitzer et al. 2008; Stadler et al. 2001). Ju and Rogers
(1996) used stromatal pigments to characterize Hypoxylon,
while Stadler et al. (2014) used a polyphasic taxonomic approach in their world monograph of Daldinia.
Dennis (1961) was the first author to attempt the segregation of the Xylariaceae into three subfamilies,
“Hypoxyloideae”, “Thamnomycetideae” and “Xylarioideae”.
However, this segregation was still based on generic concepts
that heavily relied on stromatal macromorphology,
disregarding the asexual states, and does in no way relate to
the modern concepts of the family. In addition, the subfamilies
were not validly published (cf. Bitzer et al. 2008), and the
“Thamnomycetideae” sensu Dennis later proved untenable
Fungal Diversity
Fig. 35 Vialaea mangiferae (holotype). a Herbarium specimen. b
Appearance of ascomata on substrate. c Horizontal cross section of
ascomata. d Vertical cross section of ascomata. e Peridium. f
Paraphyses. g J+, subapical apparatus of ascus. h–j Asci. k–n
Ascospores. o Surface view of culture on MEA. p Reverse view of
culture on MEA. q Conidiomata. r Conidia attached to conidiogenous
cells. s Conidia. Scale bars: b=200 μm, c=100 μm, d=200 μm, f, g=
50 μm, h, n=20 μm, q=1 mm, r, s =10 μm
as it was found out by a polyphasic approach that the affinities
of Thamnomyces s. str. are with Daldinia and allies whereas
other species of Thamnomyces are related to Xylaria (Stadler
et al. 2010b) . Nevertheless, some authors have used the remaining “subfamilies” in recent work, discriminating the taxa
with “nodulisporium-like anamorphs” sensu Ju and Rogers
(1996) (i.e., the hypoxyloid Xylariaceae) from those with
“geniculosporium-like anamorphs” (i.e., Xylaria and allies).
These subfamilies have been used in subsequent studies
(Daranagama et al. 2014, 2015; Hsieh et al. 2005, 2010;
Stadler et al. 2010a, c, 2013; Tang et al. 2009). Several
polythetic and molecular phylogenetic studies (Daranagama
et al. 2015; Maharachchikumbura et al. 2015; Stadler et al.
2010c) have indicated the possible need for the separation of
“Hypoxyloideae” and “Xylarioideae” as separate families.
However, molecular data are so far only available for a small
fraction of the existing species and no reliable data are available for several genera. Therefore, the above classification is
widely based on a combination of molecular data and morphological features relating to the type of asexual morph (cf.
Stadler et al. 2013).
Xylaria Hill ex Schrank, Baier. Fl. (München) 1: 200
(1789).
Facesoffungi number: FoF00696
Saprobic on dead wood. Sexual morph: Stromata extremely variable in size and shape, simple to branched from
Fungal Diversity
the base, nearly sessile or arising from long rooted stipes,
whitish when immature, gradually turning silvery grey and
eventually black, longitudinally furrowed delimiting narrow
strips, perithecial contours most often inconspicuous in welldeveloped stromata, leathery, black, interior solid, homogeneous, white to cream coloured, with a slightly darker core,
stipes cylindrical to strap-like, black, with a hairy-tomentose
broadened base. Ascomata subglobose, immersed to slightly
exposed. Ostioles raised-discoid, grey brown to black, often
with a low conical papilla at the center. Hamathecium comprising sparse, hypha-like, hyaline, septate paraphyses. Asci
unitunicate, (6–) 8-spored, cylindrical, long pedicellate, usually with a J+, tubular, apical apparatus. Ascospores overlapping uniseriate in the ascus, ellipsoid-inequilateral, with narrowly to broadly rounded ends, medium brown, smoothwalled, some with a fugacious cellular appendage usually
disappearing at maturity, usually with two large guttules, with
a conspicuous straight germ slit, 1/2–4/5 of spore-length on
the flattened side. Asexual morph: Geniculosporium or
geniculosporium-like. (Ju and Rogers 1996; Stadler et al.
2014).
Type species: Xylaria hypoxylon (L.) Grev., Fl. Edin.: 355
(1824) (Fournier et al. 2011; Persoh et al. 2009; Stadler et al.
2014)
= Clavaria hypoxylon L., Sp. pl. 2: 1182 (1753)
Notes: Xylaria is the largest and the type genus in the family Xylariaceae with more than 700 epithets listed in Index
Fungorum (2015). Most Xylaria species are characterised by
massive stromata which can varied in colour, shape and size
based on the location, cylindrical asci with apical apparatus
and dark ascospores unique to the family as well as their
geniculosporium-like asexual morphs (Ju and Rogers 1996;
Stadler et al. 2013). Stromata of most of the known Xylaria
species occur on decayed dicotyledonous wood, fewer on
monocotyledons, fruits, seeds and fallen leaves.
Interestingly, a number of Xylaria species that comprise the
subgenus Pseudoxylaria are associated with termite nests
mainly of macrotermitine termites, rather than plants. They
form a monophyletic clade in an otherwise polyphyletic
mega-genus (Hsieh et al. 2010).
Hypoxyloideae
Hypoxyloideae or hypoxyloid Xylariaceae have
nodulisporium–like asexual morphs (Ju and Rogers 1996).
The genus Nodulisporium was introduced by Preuss (1849),
and is typified by Nodulisporium ochraceum Preuss. Stadler
et al. (2013) stated that N. ochraceum may be associated with
Hypoxylon howeanum Peck. There are 27 generic names
listed in Stadler et al. (2013) which have been included in
the subfamily Hypoxyloideae.
Xylarioideae
The subfamily Xylarioideae has conidiogenous structures
resembling those of Geniculosporium (Ju and Rogers 1996;
Stadler et al. 2013). Chesters and Greenhalgh (1964)
introduced Geniculosporium with Geniculosporium serpens
Chesters & Greenh. as the type species (whch is actually the
asexual state of Nemania serpens. hence the name is now
obsolete). Stadler et al. (2013) have listed 38 asexual and
sexual genera that can be included in the subfamily
Xylarioideae.
The phylogeny of the Xylariaceae is currently being examined in a separate study, using morphological and chemotaxonomic data and a multi-gene genealogy. Therefore, no detailed information is being provided in the current paper.
Acknowledgments S. Maharachchikumbura thanks the Featured microbial resources and diversity investigation in Southwest Karst area
(2014FY120100) for funding. The authors extend their sincere appreciations to the Deanship of Scientific Research at King Saud University for
its funding this Prolific Research Group (PRG-1436-09). Kevin D. Hyde
thanks the Chinese Academy of Sciences, project number 2013T2S0030,
for the award of Visiting Professorship for Senior International Scientists
at Kunming Institute of Botany. Y.P. Xiao and T.C. Wen are grateful to
The National Natural Science Foundation of China (No.31460012 &
No.31200016). Shi-Ke Huang and J.C. Kang are grateful to the
Agricultural Science and Technology Foundation of Guizhou Province
Nos. NY [2013]3042), the International Collaboration Plan of Guizhou
Province (No. G [2012]7006) and the Innovation Team Construction for
Science And Technology of Guizhou Province (No. [2012]4007) from
the Science and Technology Department of Guizhou Province, China.
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