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Phylogenetic relationships and morphological reappraisal of Melanommataceae (Pleosporales)

October 2015
Fungal Diversity 74(1)

October 2015
74(1)

DOI:10.1007/s13225-015-0350-9

Authors:

Qing Tian

Mae Fah Luang University

Jian-Kui Jack Liu

University of Electronic Science and Technology of China

Kevin David Hyde

Mae Fah Luang University

Dhanushka N. Wanasinghe

Kunming Institute of Botany (CAS)

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The family Melanommataceae is widespread in temperate and subtropical regions and species invariably occur on twigs or bark of various woody plants in terrestrial, marine or freshwater habitats. In this paper, the type species of 26 genera of the family are re-described and illustrated. A multi-gene phylogeny based on maximum likelihood and Bayesian analyses of LSU, SSU, RPB2 and EF-1α sequence data of species of Melanommataceae is provided. The new genera, Muriformistrickeria, Pseudostrickeria and Thysanolaenae are introduced. Anomalemma is synonymized under Exosporiella. Acrocordiopsis, Astrosphaeriella, Beverwykella, Caryosporella, Sporidesmiella and Pseudotrichia are excluded from Melanommataceae based on molecular phylogenetic analyses. Presently, 20 genera are accepted in Melanommataceae. Based on the phylogenetic data, five new species, Byssosphaeria siamensis, Herpotrichia vaginatispora, Pseudostrickeria muriformis, Pseudostrickeria ononidis and Muriformistrickeria rubi, are introduced.

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Phylogenetic relationships and morphological reappraisal

of Melanommataceae (Pleosporales)

Qing Tian

1,2,3,4,5

& Jian Kui Liu

4,5,6

& Kevin D. Hyde

1,2,3,4,5,11

Dhanushka N. Wanasinghe

1,2,3,4,5

& Saranyaphat Boonmee

4,5

& Subashini C. Jayasiri

4,5

Zong Long Luo

4,5

& Joanne E. Taylor

& Alan J. L. Phillips

& Darbhe J. Bhat

9,10

Wen Jing Li

1,2,3,4,5

& Hiran Ariyawansa

4,5,6

& Kasun M. Thambugala

4,5,6

E. B. Gareth Jones

& Putarak Chomnunti

4,5

& Ali H. Bahkali

& Jian Chu Xu

1,2,3

Erio Camporesi

12,13,14

Received: 25 September 2015 /Accepted: 19 October 2015

School of Science 2015

Abstract The family Melanommataceae is widespread in tem-

perate and subtropical regions and species invariably occur on

twigs or bark of various woody plants in terrestrial, marine or

freshwater habitats. In this paper, the type species of 26 genera

of the family are re-described and illustrated. A multi-gene

phylogeny based on ma ximum likelihood and Bayesian

analyses of LSU, SSU, RPB2 and EF-1α sequence data of

species of Melanommataceae is provided. The new genera,

Muriformistrickeria, Pseudostrickeria and Thysanolaenae

are introduced . Anomalemma is synonymized under

Exosporiella. Acr ocordiopsis, Astrosphaeriella, Beverwykella,

Caryosporella, Sporidesmiella and Pseudotrichia are ex-

cluded from Melanommataceae basedonmolecularphy-

logenetic analyses. Presently, 20 genera are accepted in

Melanommataceae. Based on the phylogenetic data, five new

species, Byssosphaeria siamensis, Herpotrichia vaginatispora,

Pseudostrickeria muriformis, Pseudostrickeria ononidis and

Muriformistrickeria rubi, are introduced.

Keywords Byssosphaeria

Herpotrichia

Multi-gene

Phylogeny

Muriformistrickeria

New species

Pseudostrickeria

Taxonomy

Thysanolaenae

Type

Introduction

The Pleosporales is the largest order in the class

Dothideomycetes (Kirk et al. 2008; Zhang et al. 2012; Hyde

* Kevin D. Hyde

kdhyde3@gmail.com

Key Laboratory for Plant Diversity and Biogeography of East Asia,

Kunming Institute of Botany, Chinese Academy of Sciences,

Kunming 650201, Yunnan, People’s Republic of China

World Agroforestry Centre, East and Central Asia,

Kunming 650201, Yunnan, People’s Republic of China

Center for Mountain Ecosystem Studies, Kunming Institute of

Botany, Chinese Academy of Sciences, Kunming 650201,

Yunnan, China

School of Science, Mae Fah Luang University, Chiang Rai 57100,

Thailand

Center of Excellence in Fungal Research, Mae Fah Luang University,

Chiang Rai 57100, Thailand

Guizhou Key Laboratory of Agricultural Biotechnology,

Guizhou Academy of Agricultural Sciences, Guiyang 550006,

Guizhou, China

Royal Botanic Garden Edinburgh, 20A Inverleith Row,

Edinburgh EH3 5LR, UK

UCIBIO, Faculdade de Ciências e Tecnologia, Universidade Nova de

Lisboa, 2829-516 Caparica, Portugal

Formerly, Department of Botany, Goa University, Goa-403206, India

Res: No. 128/1–J, Azad Housing Society, Curca,

P. O. Goa Velha-403108, India

Department of Botany and Microbiology, College of Science,

King Saud University, P.O. Box: 2455, Riyadh 1145, Saudi Arabia

A.M.B. Gruppo Micologico Forlivese BAntonio Cicognani^,

Via Roma 18, Forlì, Italy

A.M.B. Circolo Micologico BGiovanni Carini^,

C.P. 314 Brescia, Italy

Società per gli Studi Naturalistici della Romagna,

C.P. 144 Bagnacavallo, RA, Italy

Fungal Diversity

DOI 10.1007/s13225-015-0350-9

et al. 2013; Wijayawardene et al. 2014). Intensive sampling

and multigene phylogenetic analyses have placed

Melanommatales as a synonym of Pleosporales and 40 fam-

ilies were included in the order Pleosporales (Liew et al.

2000; Lumbsch and Lindemuth 2001;Zhangetal.2012;

Hyde et al. 2013; Wijayawardene et al. 2014).

Melanommataceae is an interesting family of the

Pleosporales, but it is polyphyletic and in need of revision

(Kirk et al. 2008). Most species of this family are saprobic

or hyperparasitic and occur mainly on twigs or bark of various

woody plants in terrestrial, marine or freshwater habitats, and

are widespread in temperate and subtropical regions (Zhang

et al. 2012;Hydeetal.2013).

The family Me lanommataceae was est ablished by

Winter (1885) and is characterized by globose or de-

pressed perithecial ascomata, bitunicate and fissitunicate

asci, pigmented and phragmosporous ascospores (Barr

1990a; Sivanesan 1984; Zhang et al. 2012;Hydeetal.

2013). The key diagnostic character of the species in

Melanommataceae is the trabec ulate pseudo paraphyses

(Sivanesan 1984;Barr1990a;Liewetal.2000;Kirk

et al. 2001). As Barr (198 3) cons idered trabe culate

pseudoparaphyses to be important at the ordinal level,

the order Melanommatales was introduced to accommo-

date all taxa with trabeculate pseudoparaphyses

(Sporormia-type ce ntrum developmen t), which can be

distinguished from cellular pseudoparaphyses

(Pleospora-type centrum development) in Pleosporales.

The family Melanommataceae was reviewed by Barr

(1990a) who included Ostropella, Keissleriella,

Strickeria, Byssosphaeria and Melanomma . These genera

are saprobic, especially on woody substrates, or occa-

sionally hyperparasitic on ascomycetes, e.g.,

Keissleriella Höhn. However, recent phylogenetic analy-

ses did not support this cla ssification a nd instead has

shown t hat Melanom matales should be assigned to

Pleosporales (Berbee 1996;Liewetal.2000; Lumbsch

and Lindemuth 2001;Schochetal.2006; Kruys et al.

2006;Wangetal.2007). Consequently, Eriksson (2006)

synonymized Melanommatales under Pleosporales.

Following the phylogenetic studies by Zhang et al.

( 2009b, 2012), Keissleriella was p laced in the family

Lentitheciaceae. von Höhnel (1918)andBarr(1990a)

proposed that Xenolophium should be a synonym of

Ostropella. However, based on the differences in their

ascomatal surface structure, morphology of the ascomatal wall

in longitudinal section and ascospore morphology, Huhndorf

(1993)maintainedOstropella and Xenolophium as separate

genera in Melanommataceae. Based on LSU sequence data,

Mugambi and Huhndorf (2009)demonstratedthatOstropella

albocincta clustered together with Xenolophium applanatum

in the family Platystomaceae, thus, Zhang et al. (2012)for-

mally excluded it from M elanommataceae despite the

trabeculate pseud oparaphyses. Lumbs ch and Huhndorf

(2010) accepted 14 genera and some genera were excluded,

viz. Bimuria, Ostropella, Trematosphaeria and Xenolophium

(Mugambi and Huhndorf 2009; Suetrong et al. 2011).

Between 1990 and 2014, Melanommataceae was revised

with additions and exclusions of genera (Table 1). Lumbsch

and Huhndorf (2007) accepted 18 genera of which six were

questionably placed. Kirk et al. (2008)accepted21genera,

whereas, Zhang et al. (2012) accepted nine genera with another

four questionably placed. Wijayaward ene et al. (2014)included

the sexual and asexual morphs and listed 18 genera. There have

been several reports of asexual morphs of melanomma taceous

genera, although these need to be confirmed in the light of

modern taxonomic treatments and molecular data. The reported

asexual morphs of Melanommataceae are coelomycetous and

hyphomycetous. Herpotrichia is reported as having a

Pyrenochaeta asexual morph with or without pycnidial setae

(Sivanesan 1984). Aposphaeria and Pho ma-like asexual

morphs have been reported for Melanomma species

(Chesters 1 938; Sivanesan 1984). Similarly, the asexual

morphs of Karstenula are reported as coelomycetous (i.e.,

Microdiplodia, Constantinescu 1993). The asexual morph of

Exosporiella was reported as Anomalemma (Sivanesan 1983),

and that of Byssosphaeria as Pyrenochaeta (Barr 1984).

Ohleria brasiliensis Starbäck has been linked with

Monodictys putredinis (Wallr.) S. Hughes (Samuels 1980).

There have been several recent studies showing the poly-

phyletic nature of the family Melanommataceae (Liew et al.

2000; Kodsueb et al. 2006; Kruys et al. 2006;Wangetal.

2007; Mugambi and Huhndorf 2009; Zhang e t al. 2009b,

2012; Hyde et al. 2013;Wijayawardeneetal.2014; Liu et al.

2015). The genera Byssosphaeria and Melanomma are, how-

ever, strongly supported within the family Melanommataceae.

Molecular studies showed Pleomassariaceae to be a synonym

of Melanommataceae (Zhang et al. 2009b; Tanaka et al. 2010).

This is because the types of Melanomma (M. pulvis-pyrius)

and Pleomassaria (

P. siparia) clustered in the same clade with

strong bootstrap support. Subsequently, Zhang et al. (2012)

reinstated Pleomassariaceae as a separate family in the order

Pleosporales based on Melanommataceae being a well-

supported monophyletic clade.

Recently, numerous genera and species have been accepted

in Melanommataceae based on phylogenetic studies which

confirmed the placement of several taxa (Liu et al. 2015;Su

et al. 2015). Many taxa are asexual morphs that can-not be

linked to the sexual morph in Melanommataceae. Therefore,

phylogenetic analysis are essential to confirm the conection

between sexual and asexual morphs within the

Melanommataceae. In the present study, we accept 13 sexual

morph genera viz. Asymmetricospora, Bertiella, Bicrouania,

Byssosphaeria, Calyptronectria, Herpotrichia,

Mamillisphaeria, Melanomma, Muriformistrickeria,

Navicella, Ohleria, Pseudostrickeria, Sarimanas and seven

Fungal Diversity

asexual morph genera in Melanommataceae,viz.

Aposphaeria, Exos poriella, Mycopappus, Nigrolentilocus,

Monotosporella, Phragmocephala and Xenostigmina.The

aim of this study was to reappraise the placement of several

genera in Melanommataceae based on a phylogenetic analysis

and on morphological characteristics, and to provide a back-

bone tree for this group of taxa.

Materials and methods

Herbarium examination

Herbarium specimens, including type specimens were loaned

from the following herbaria: BRIP, IFRD, G, K, NYand S (see

http://sweetgum.nybg.org/ih/index.php for full names ).

Table 1 Various treatments of genera in Melanommataceae

Barr (1990a) Lumbsch and Huhndorf (2007) Kirk et al. (2008) Lumbsch and Huhndorf

(2010)

Hyde et al. (2011) (Asexual morph

included)

Byssosphaeria Acrocordiopsis Anomalemma Acrocordiopsis Aposphaeria

Keissleriella ?Anomalemma Asymmetricospora ?Anomalemma Beverwykella

Melanomma ?Astrosphaeriella Byssosphaeria Asymmetricospora Nigrolentilocus (Sexual=Melanomma)

Ostropella Asymmetricospora Calyptronectria Bertiella Monodictys (Sexual=Ohleria)

Strickeria Bicrouania Caryosporella Bicrouania Monotosporella

?Bimuria Dangeardiella Byssosphaeria Pyrenochaeta (Sexual=Byssosphaeria)

Byssosphaeria Javaria Calyptronectria Sporidesmiella

Calyptronectria Karstenula ?Caryosporella

?Caryosporella Melanomma Herpotrichia

Javaria Mycopepon Javaria

Karstenula Ohleria ?Mamillisphaeria

?Mamillisphaeria Ostropella Melanomma

Melanomma Xenolophium Ohleria

Ohleria Pseudotrichia

Ostropella

Pseudotrichia

?Trematosphaeria

Xenolophium

Zhang et al. (2012) Wijayawardene et al. (2012)

(Asexual morph included)

Hyde et al. (2013) Wijayawardene et al.

(2014)

This study

Anomalemma Aposphaeria Anomalemma Aposphaeria Aposphaeria

Astrosphaeriella Beverwykella Asymmetricospora Asymmetricospora Asymmetricospora

Bertiella

?Asymmetricospora Exosporiella (Sexual=Anomalemma) Bertiella Bertiella Bicrouania

Bertiella Nigrolentilocus (Sexual=Melanomma) Beverwykella Beverwykella Byssosphaeria

?Bicrouania Monodictys (Sexual=Ohleria) Bicrouania Bicrouania Calyptronectria

Byssosphaeria Monotosporella Byssosphaeria Byssosphaeria Exosporiella

Calyptronectria Pyrenochaeta (Sexual=Byssosphaeria) Calyptronectria Calyptronectria Herpotrichia

?Caryosporella Sporidesmiella Caryosporella Caryosporella Mamillisphaeria

Herpotrichia Exosporiella Exosporiella Melanomma

?Mamillisphaeria Herpotrichia Herpotrichia Monotosporella

Melanomma Mamillisphaeria Mamillisphaeria Muriformistrickeria

Ohleria Melanomma Melanomma Mycopappus

Pseudotrichia Nigrolentilocus Mycopappus Navicella

Ohleria Navicella Nigrolentilocus

Pseudotrichia Nigrolentilocus Ohleria

Sporidesmiella Ohleria Pseudostrickeria

Sporidesmiella Phragmocephala

Xenostigmina Sarimanas

Xenostigmina

Fungal Diversity

Ascomata were rehydrated in 3 % KOH or water and stained

with cotton blue prior to examination and sectioning. India ink

was added to water mounts to visualize gelatinous sheaths and

ascospore appendages. Cotton blue and Congo red stains were

used to verify the septation of ascospores and internal

elements of the hamatheciu m. Permanent slide s were

prepared by mounting material in lactoglycerol and sealing

the cover slips with clear nail polish. Ascomata were

examined with a stereo microscope (Olympus SZH10, free

hand sections of the fruiting structures were cut with a razor

blade and mounted in water. Squash mounts of ascomata were

prepared in a droplet of distilled water on a clean slide.

Micromorphological characters were determined with a

Nikon ECLIPSE 80i compound mic roscope and images

were captured with a C anon EOS 550D digita l camera.

Differential interference contrast microscopy was used for

hyaline structures. Measurements were made with Tarosoft

(R) Image FrameWork version 0.9.7. Photographic plates

were prepared in Adobe Photoshop version CS3 (Adobe

Systems Inc., The United States).

Terminologies mainly follow Ulloa and Hanlin (2000)and

Zhang et al. (2009b, 2012). In addition, ascomata size is de-

fined as small: <300μm diam., medium: from 300 to 600μm

diam., large: >600μm diam. In this study, a question mark

before a genus name, indicates an uncert ain placement in

Melanommataceae.

Fungal isolation, morphology and asexual state study

Fresh samples of Melanommataceae were collected from dif-

ferent sites in Italy and Thailand and brought to the laboratory

in Zip lock plastic bags. Collected specimens were incubated

in a moist chamber for 1–2 days at 25 °C. After morphological

examination, single ascospore isolations were made following

the method described by Chomnunti et al. (2014).

Germinating ascospores were transferred aseptically to malt

extract agar (MEA) and potato dextrose agar (PDA) plates and

grown at 16 °C in daylight. Sporulation was induced on water

agar (WA) plates with sterile pine needles (Phookamsak et al.

2015). Colony colour and other characters were observed and

recorded after 1 week and again after 3 weeks.

Specimens are deposited in the herbarium of Kunming

Insititute of Botany, Chinese Academy of Sciences (HKAS),

Kunming, China and Mae Fah Luang University (MFLU),

Chiang Rai, Thailand. Living cultures are deposited in Mae

Fah Luang University Culture Collection (MFLUCC), and

duplicated in the International Collection of Microorganisms

from Plants (ICMP), Landcare Research, New Zealand and

Kunming Insititute of Botany, Chinese Academy of Sciences

(KUMCC), Kunming, China. Taxa were registered in Index

Fungorum (Kirk et al. 2015)(www.indexfungorum.org/

names/names.asp) and Facesoffungi (Jayasiri et al. (2015)).

Fungal DNA extraction, PCR amplification, sequencing

and sequence alignment

Biospin Fungus Genomic DNA Extraction Kit-BSC14S1

(BioFlux®, P.R. China) was used to extract DNA from fresh

mycelium grown on PDA at 16 °C for 4 weeks using the

instructions of the manufacturer (Hangzhou, P.R. China).

Phylogenetic analyses were based on partial sequences of

four loci, small subunit rDNA (SSU), large subunit (LSU),

translation elongation factor 1-alpha gene (TEF 1α) and sec-

ond largest subunit of RNA polymerase II (RPB2). LSU re-

gion was amplified using the primers LROR and LR5

(Vilgalys and Hester 1990), SSU was amplifie d using the

primers NS1 and NS4 (White et al. 1990), TEF 1α was am-

plified using primers EF1-983F and EF1-2218R ( Rehner

2001) and RPB2 was amplified using the primers fRPB2-5F

and fRPB2-7cR (Liu et al. 1999).

Polymerase chain reaction (PCR) was carried out using the

following protocol: The final volume of the PCR reaction was

25 μL and contained 12.5 μLof2×PowerTaqPCR

MasterMix (a premix and ready to use solution, including

0.1 Units/μl Taq DNA Polymerase, 500 μ

mdNTPMixture

each (dATP, dCTP, dGTP, dTTP), 20 mM Tris–HCl pH 8.3,

100 Mm KCl, 3 mM MgCl

, stabilizer and enhancer), 1 μLof

each primer (10 μM), 1 μL genomic DNA extract and 9.5 μL

deionised water. The reaction was then allowed to run for

35 cycles. The annealing temperature was 55 °C for LSU,

SSU & TEF 1α and initially denaturation 94 °C for 3 min,

denaturation at 94 °C for 30 s, annealing for 50 s, elongation at

72 °C for 1 min, and final extension at 72 °C for 10 min. The

PCR thermal cycle program for the partial RNA polymerase

second largest subunit (RPB2) was followed as initially 95 °C

for 5 min., followed by 40 cycle of denaturation at 95 °C for

1 min, annealing at 52 °C for 2 min., elongation at 72 °C for

90 s, and final extension at 72 °C for 10 min. PCR amplifica-

tionwasconfirmedon1%agarose electro phores is gels

stained with ethidium bromide. The amplified PCR fragments

were sequenced by BGI, Ltd Shenzhen, P.R. China).

Sequence data are deposited in GenBank (Table 2).

Phylogenetic analyses

The closest taxa to our strains were determined with standard

nucleotide blast searches in GenBank (http://www.ncbi.nlm.

nih.gov/), and sequences of representative species were

selected as in Su et al. (2015).

Combined analysis of LSU, SSU, RPB2 and EF-1α se-

quence data of closest relatives in Aigialaceae,

Bambusicolaceae, Lentitheciaceae, Massarinaceae,

Melanommataceae , Montag nulaceae, Morosphaeriaceae,

Pleomassariaceae, Roussoellaceae, Thyr idariaceae and

Trematosphaeriaceae were used to confirm the phylogenetic

placement of Melanommataceae in Pleosporales. Hysterium

Fungal Diversity

Table 2 GenBank and culture collection accession numbers of species treated in the phylogenetic study. Newly generated sequences are shown in bold

Species Culture collection GenBank accession numbers

LSU SSU EF1-α RPB2

Aposphaeria corallinolutea CBS 131287 JF740330 / / /

Aposphaeria corallinolutea CBS 131286 JF740329 / / /

Aposphaeria populina CBS 543.70 EU754130 EU754031 / /

Aposphaeria populina CBS 350.82 JF740265 / / /

Beverwykella pulmonaria CBS 283.53 GU301804 / / GU371768

Beverwykella pulmonaria MFUCC 15–0183 KT934251 ///

Bertiella macrospora IL 5005 GU385150 / / /

Byssosphaeria jamaicana SMH1403 GU385152 / GU327746 /

Byssosphaeria jamaicana SMH3085 GU385154 / / /

Byssosphaeria rhodomphala SMH3086 GU385155 / / /

Byssosphaeria rhodomphala GKM L153N GU385157 / GU327747 /

Byssosphaeria salebrosa SMH2387 GU385162 / GU327748 /

Byssosphaeria schiedermayeriana SMH3157 GU385163 / GU327745 /

Byssosphaeria schiedermayeriana GKM 152 N GU385168 / GU327749 /

Byssosphaeria villosa GKM 204 N GU385151 / GU327751 /

Byssosphaeria musae MFLUCC 11–0146 KP744477 KP753947 / /

Byssosphaeria siamensis MFLUCC 10–0099 KT289895 KT289897

Byssosphaeria schiedermayeriana MFLUCC 10–0100 KT289894 KT289896

Caryosporella rhizophorae JK 5302A GU479784 / / /

Herpotrichia diffusa CBS 250.62 DQ678071 DQ678019 DQ677915 DQ677968

Herpotrichia juniperi CBS 200.31 DQ678080 DQ678029 DQ677925 DQ677978

Herpotrichia macrotricha GKM 196 N GU385176 / GU327755 /

Herpotrichia macrotricha SMH 269 GU385177 / GU327756 /

Herpotrichia parasitica CBS451.73 GQ387617 GQ387556 / /

Herpotrichia vaginatispora MFLUCC 13–0865 KT934252 KT934256 KT934260 /

Melanomma pulvis-pyrius CBS 124080 GU456323 GU456302 GU456265 GU456350

Melanomma pulvis-pyrius CBS 109.77 FJ201986 FJ201987 GU456274 GU456359

Melanomma pulvis-pyrius CBS 371.75 FJ201988 FJ201989 GU349019 GU371798

Melanomma rhododendri ANM 73 GU385198 / / /

Muriformistrickeria rubi MFLUCC 15–0681 KT934253 KT934257 KT934261 /

Sarimanas shirakamiense MAFF 242969 LC001716 LC001713 / /

Sarimanas shirakamiense MAFF 244768 LC001715 LC001712 / /

Sarimanas pseudofluviatile MAFF 239465 LC001714 LC001711 / /

Sporormiella minima CBS 524.50 DQ678056 DQ678003 DQ677897 DQ677950

Sporidesmiella fusiformis HKUCC 10831 DQ408577 / / DQ435079

Pseudostrickeria muriformis MFLUCC 13–0764 KT934254 KT934258 KT934262 /

Pseudostrickeria ononidis MFLUCC 14–0949 KT934255 KT934259 KT934263 KT934264

Pseudotrichia mutabilis

SMH 1541 GU385209 / / /

Pseudotrichia mutabilis SMH 5288 GU385210 / / /

Pseudotrichia thailandica MFLUCC 1 1–0155 KP744510 KP753968 / /

Pseudotrichia rubriostiolata MFLUCC 11–0138 / KP744508 / /

Pseudotrichia rubriostiolata MFLUCC 11–0156 KP744509 KP753967 / /

Pseudotrichia rubriostiolata MFLUCC 11–0137 KP744507 KP753966 / /

Pseudotrichia guatopoensis SMH 4535 GU385202 / GU327774 /

Phragmocephala atra MFLUCC 15–0021 KP698729 KP698725 / /

Phragmocephala garethjonesii MFLUCC 15–0018 KP698730 KP698726 / /

Xenostigmina zilleri CBS 115686 GU253858 / / /

Fungal Diversity

Table 2 (continued)

Species Culture collection GenBank accession numbers

LSU SSU EF1-α RPB2

Xenostigmina zilleri CBS 115685 GU253857 / / /

Xenostigmina zilleri CBS 124108 FJ839675 / / /

Monotosporella tuberculata CBS 256.84 GU301851 / GU349006 /

Mycopappus aceris CBS 124109 FJ839660 / / /

Prosthemium betulinum VM20040721 AB553754 AB553644 / /

Pleomassaria siparia CBS 279.74 DQ678078 DQ678027 DQ677923 DQ677976

Roussoella nitidula MFLUCC 11–0634 KJ474842 / KJ474851 KJ474858

Roussoella nitidula MFLUCC 11–0182 KJ474843 / KJ474852 KJ474859

Roussoella hysterioides CBS 125434 AB524622 AB524481 AB539115 AB539102

Phaeodothis winteri CBS 182.58 GU301857 GU296183 / /

Bimuria novae-zelandiae CBS 107.79 AY016356 AY016338 DQ471087 DQ470917

Kalmusia scabrispora KT 2202 AB524594 AB524453 AB539107 AB539094

Karstenula rhodostoma CBS 690.94 GU301821 GU296154 GU349067 GU371788

Paraphaeosphaeria michotii CBS 591.73 GU456326 GU456305 GU456267 GU456352

Montagnula opulenta CBS 168.34 DQ678086 / / DQ677984

Massarina cisti CBS 266.62 FJ795447 FJ795490 / FJ795464

Massarina eburnea CBS 473.64 GU301840 GU296170 GU349040 GU371732

Byssothecium circinans CBS 675.92 AY016357 AY016339 GU349061 DQ767646

Lentithecium aquaticum CBS 123099 GU301823 GU296156 GU349068 GU371789

Lentithecium fluviatile CBS 122367 GU301825 GU296158 GU349074 /

Katumotoa bambusicola MAFF 239641 AB524595 AB524454 AB539108 AB539095

Keissleriella cladophila CBS 104.55 GU301822 GU296155 GU349043 GU371735

Morosphaeria ramunculicola JK 5304B GU479794 GU479760 / GU479831

Helicascus nypae BCC 36752 GU479789 GU479755 GU479855 GU479827

Trematosphaeria pertusa CBS 122368 FJ201990 FJ201991 KF015701 FJ795476

Trematosphaeria pertusa CBS 122371 GU301876 FJ201993 KF015702 GU371801

Aigialus grandis BCC 18419 GU479774 GU479738 GU479838 GU479813

Aigialus mangrovis BCC 33564 GU479777 GU479742 GU479841 GU479816

Astrosphaeriella aggregata KT 767 AB524590 AB524449 / /

Astrosphaeriella aggregata KT 984 AB524591 AB524450 AB539105 AB539092

Astrosphaeriella bakeriana CBS 115556 GU301801 / GU349015 GU357752

Astrosphaeriella stellata MFLUCC10-0555 JN846723 JN846733 / /

Astrosphaeriella stellata MFLUCC10-0095 JN846720 JN846741 / /

Bambusicola massarinia MFLUCC 11–0389 JX442037 JX442041 / /

Bambusicola splendida MFLUCC 11–0439 JX442038 JX442042 / /

Thyridaria rubronotata CBS 419.85 GU301875 / GU349002 GU371728

Massariosphaeria phaeospora CBS 611.86 GU301843 GU296173 / GU371794

Acrocordiopsis patilii BCC 28166 GU479772 GU479736 / GU479811

Acrocordiopsis patilii BCC 28167 GU479773 GU479737 / GU479812

Salsuginea ramicola KT 2597.1 GU479800 GU479767 GU479861 GU479833

Hysterium angustatum CBS 123334 FJ161207 FJ161167 FJ161 1 1 1 FJ161129

Hysterium angustatum CBS 236.34 FJ161180. GU397359 FJ161096 FJ161117

ANM A.N. Miller, ATCC

American Type Culture Collection Virginia, USA, BCC BIOTEC Culture Collection, Bangkok, Thailand, CBS Centraalbureau

voor Schimmelcultures, Utrecht, The Netherlands, GKM G.K. Mugambi, HKUCC University of Hong Kong Culture Collection, Department of Ecology

and Biodiversity, Hong Kong, China, IL I. Lopez, JK J. Kohlmeyer, KT K. Tanaka, MAFF Ministry of Agriculture, Forestry and Fisheries, Japan,

MFLUCC Mae Fah Luang University Culture Collection, ChiangRai, Thailand, SMH S.M. Huhndorf, VM V.A. Mel'nik

Fungal Diversity

angustatum Alb. & Schwein (CBS 123334) and

H. angustatum Alb. & Schwein (CBS 236.34) was selected

as the outgroup taxa (Fig. 1).

The other sequences used in the analyses (Table 2)were

obtained from GenBank. The multiple alignments were made

by MAFFT v. 7.036 (Katoh and Standley 2013), and adjusted

manually for improvement where necessary using Mega 6.0.5

(Tamura et al. 2013), BioEdit v. 7.2 (Hall 1999) and ClustalX

v. 1.83 (Thompson et al. 1997).

MODELTEST v. 2.0 (Nylander 2004) following Akaike

Information Criterion was used to determine the best-fit model

of evolution for each data set for Bayesian and Maximum

Likelihood analyses.

Maximum-likelihood (ML) analysis was performed in

RAxML (Stamatakis et al. 2008) implemented in raxmlGUI

v.0.9b2 (Silvestro and Michalak 2011). One thousand non-

parametric bootstrap iterations were employed with the avail-

able models of generalized time reversible (GTRGAMMAI

model.) and a discrete gamma distribution (Stamatakis et al.

2008; Liu et al. 2011). Maximum likelihood bootstrap values

equal or greater than 50 % are given as the first set of numbers

above the nodes (Fig. 1).

A Bayesian analysis was conducted with MrBayes v. 3.1.2

(Huelsenbeck and Ronquist 2001) to evaluate Posterior prob-

abilities (PP) (Rann ala and Yang 1996; Zhaxybayeva and

Gogarten 2002) by Markov Chain Monte Carlo sampling

(BMCMC). Six simultaneous Markov chains were run for 1,

000,000 generations and trees were sampled every 100th gen-

eration and 10,000 trees were obtained. The first 2000 trees,

representing the burn-in phase of the analyses, were discarded,

while the remaining 8000 trees were used for calculating pos-

terior probabilities in the majority rule consensus tree (critical

value for the topological convergence diagnostic is 0.01) (Cai

et al. 2006). Bayesian Posterior Probabilities (BYPP) equal or

greater than 0.90 is given below or above each node (Fig. 1).

Results

Phylogenetic analyses

The phylogenetic tree in Fig. 1 was constructed to vertify the

relationships of genera in Melanommataceae. The combined

LSU, SSU, RPB2 and EF-1α gene analysis comprised 89

strains with Hysterium angustatum (CBS 123334 and CBS

236.34) as the outgroup taxa. The best scoring RAxML tree

with a likelihood value is shown in Fig. 1.

Most of the core families of Pleosporales (Zhang et al.

2012;Hydeetal.2013) were included in our phylogenetic

analysis (Fig. 1). Seven strains generated in the present study

clustered well within Melanommataceae and several genera

were previously recorded in this family, such as Beverwykella,

Pseudotrichia and Pleomassaria. However, some genera

newly nested in the family Melanommataceae,suchas

Pseudostrickeria and Muriformistrickeria. Sporidesmiella lies

far away from Melanomma species and is therefore excluded

from Melanommataceae. Ten main clades could be recog-

nized within Melanommataceae (Fig. 1).

Byssosphaeria is a well-defined genus in

Melanommataceae. Our two new strains and nine reference

sequences of Byssosphaeria from previous studies (Mugambi

and Huhndorf 2009; Zhang et al. 2012; Hyde et al. 2013)

formed a well-supported clade (84 % ML BS and 1.00 BY

PP). Byssosphaeria siamense formed a c lade allied with

Byssosphaeria musae in Melanommataceae (69 % ML

BSand1.00BYPP)whichformedasistergroupwith

B. salebrosa and B. schiedermayeriana (Fuckel) M.E.

Barr with 74 % ML BS and 1.00 BY PP (Fig. 1). A

new collection of Byssosphaeria schiedermayeriana

(MFLUCC 10–0100) from Thailand, clustered with

two referenced sequences with low support. The phylo-

genetic reconstructions based on the combined datasets

analysis showed that Bertiella macrospora (IL5005)

clusteredintheByssosphaeria group (95 % ML BS

and 1.00 BY PP). Futhermore, these taxa can be distin-

guished easily on their morphological characters.

Pseudostrickeria is a new genus introduced in this study to

accommodate Pseudostrickeria muriformis and

Pseudostrickeria ononidis. They cluster together within a dis-

tinct clade in Melanommataceae (Fig. 1).

Phragmocephala and Sarimanas are newly introduced in

Melanommataceae and formed well-supported groups respec-

tively (Liu et al. 2015;Suetal.2015).

Herpotrichia has previously been placed in other families

(Kirk et al. 2008). Bar r (1984) placed Herpotrichia in

Massarinaceae, and this family was later synonymized with

Lophiostomataceae (Barr 1987). In this study, it was shown to

be polyphyletic wi th species in three clades. Herpotrichia

diffusa clustered within Byssosphaeria. Herpotrichia

macrotricha as represented by two strains obtained from

GenBank (GKM 196N and SMH 269), and formed a distinct,

strongly supported clade (Fig. 1). Herpotrichia macrotricha is

easily recognised by its distinctive broad cap-like ascomatal

apex. The new species Herpotrichia vaginatispora

(MFLUCC 13–0865) clustered in Herpotrichia forming a sis-

ter group with H. macrotricha. Herpotrichia parasitica from

GenBank formed a single clade consistently separate from the

other three species. It did not nest in Melanommataceae,while

being closely aligned with other families close to

Thyridariaceae (Fig. 1). Herpotrichia and Thyridariaceae

species share similar characters, such as immersed, erumpent

to superficial ascomata and 1- or 3-septate ascospores, while

the ascomata in Herptrichia are tomentose and usually sit on a

subiculum. Lacking of sequences data for the type species

H. rubi, thus, the natural plac ement for this genus is still

unsolved.

Fungal Diversity

Melanomma species clustered in a well-supported clade.

Thus, Melanomma pulvis-pyrius (CBS 124080, CBS 109.77

and CBS 371.75), M. rhododendri (ANM 73), Aposphaeria

populina (CBS 543.70 and CBS 350.82) and Aposphaeria

corallinolutea (CBS 131286 and CBS 131287) obtained from

GenBank formed a well-s uppor t clade. Futhermore , from

Fig. 1, it is clear that A. populina clustered together with

Melanomma pulvis-pyrius (81 % ML BS and 0.95 BY PP).

It is not possible to synonymize Aposphaeria under

Melanomma until the molecular data of the type species

Aposphaeria pulviscula is obtained.

Strains of Pseudotrichia, which has be en aligned with

Byssosphaeria and Herpotrichia, are placed in

Pleomassariaceae in the present study, along with

Pleomassaria siparia (CBS 279.74). According to Barr

(1982) Pleomassaria siparia is characterised by immersed,

depres sed, globose ascomata, containing oblong asci with

simple muriform ascospores. The ascospores are dark brown

with a verruculose surface, with 5–7transverseseptaandone

longitudinal septum in several cells. The characteristics this

species shares with others in the clad e are not obvious.

However, the placement is not clear since in our analyses,

Pseudotrichia species did not cluster together. The two new

Pseudotrichia species introduced in Liu et al. (2015)formeda

distinct clade in the phylogenetic tree, between

Roussoellaceae and Aigialaceae. The morphology of these

taxa and holotype of Pseudotrichia stromatophila

(Thambugala et al. 2014) differ greatly, therefore, a new genus

Thysanolaenae is introduced in this study to accommodate

these species.

Taxo nomy

Melanommataceae G. Winter [as ‘Melanommeae’], Rabenh.

Krypt.-Fl., Edn 2 (Leipzig) 1.2: 220 (1885)

Faces of Fungi number: FoF 01023

Saprobic or parasitic on plants in terrestrial, marine or

freshwater habitats. Sexual morph: Ascomata scattered or

gregarious, immersed to erumpent, nearly superficial, globose

to subglobose, carbonaceous or coriaceous, papillate or

epapillate. Peridium composed of heavily pigmented, thick-

walled, pseudoparenchymatous cells. Hamathecium of dense,

long, trabeculate pseudoparaphyses, embedded in a gelatinous

matrix. Asci 8-spored, bitunicate, fissitunicate, clavate to near-

ly cylindrical, pedicellate, apically rounded with an ocular

chamber. Ascospores

uniseriate or bi-seriate, fusoid to ellip-

soidal, or muriform, hyaline or brown, 1 to multi-septate, with

or without a mucilaginous sheath, wall smooth or verrucose.

Asexual morph:

Mostly coelomycetous and rarely hyphomycetous with

various conidium ontogenic structures.

Notes: In the present study, 20 genera are accepted in

Melanomm ataceae, including seven asexual morphs.

Wijayawardene et al. (2012) listed the asexual morphs as

Aposphaeria , Beverwykella, Ex osporiella, Monodi ctys,

Monotosporella, Nigrolentilocus, Pyrenochaeta and

Sporidesmiella. de Gruyter et al. (2012) showed that phyloge-

netically Ap osphaeria belongs to Melano mmataceae.

However Seifert et al. (2011) stated that Monodictys and

Monotosporella are probably heterogenous. Based on the phy-

logenetic analysis in this study, Anomalemma is synonymized

under Exosporiella. Beverwykella and Sporidesmiella are ex-

cluded from Melanommataceae (Fig. 1). Therefore, seven

asexual morphs are accepted as Aposphaeria, Exosporiella,

Monotosporella, Mycopappus, Nigrolentilocus,

Phragmocephala and Xenostigmina. Based on the mul ti-

gene phylogenetic analysis, Aposphaeria, Bertiella,

Byssosphaeria, Herpotrichia, Melanomma, Monotosporella,

Muriformistrickeria, Mycopappus, Phragmocep hala,

Pseudostrickeria, Sarimanas cluster in the

Melanommataceae. Asymmetricospora, Bicrouania,

Calyptronectria, Caryosporella, Exosporiella

Mamillisphaeria, Navicella, Nigrolentilocus and Ohleria are

retained within the Melanommataceae based on morphologi-

cal similarities, especially the coriaceous or carbonaceous

ascomata with ostioles and trabeculate pseudoparaphyses.

Key to genera in Melanommataceae

1. Sexual morph………………………………................…2

1. Asexual morph……………....................……….............14

2. Habitat freshwater or marine, saprobic…………….....….3

2. Habitat terrestrial, saprobic, parasitic or fungicolous…….4

3. Ascomata conical.......................................Mamillisphaeria

3. Ascomata globose to subglobose………......…Bicrouania

4. Ascomata nearly superficial on host, spreading on substrate

as subiculum……….…….…….…….…….…..……...........5

4. Ascomata immersed or erumpent on host……..................7

5. Ostiole with redish orange or greenish tint at

pore………………….......................................Byssosphaeria

5. Ostiole without coloured tint at pore………….....………6

6. Ascospores 1–3-septate….....………..…….....…..Bertiella

6. Ascospores 5–8-septate……………….................Navicella

7. Asci with furcate pedicel……….................…….....……..8

7. Asci without furcate pedicel…………………….....…….9

8. Ascospores 1-septate………

.…………......…Herpotrichia

8. Ascospores muriform……….....…….....…Calyptronectria

9. Ascospores lacking a mucilaginous sheath .....................10



Fig. 1 Phylogenetic tree generated by Maximum likelihood (RAxML)

analysis of combined LSU, SSU, RPB2 and EF-1α sequence data.

Bootstrap support values for maximum likelihood equal or greater than

50 % are given above the nodes. Bayesian posterior probabilities equal or

greater than 0.90 are given below the nodes. Ex-type strains and reference

strains are in bold. Newly generated sequences are in red. The tree is

rooted with Hysterium angustatum CBS 123334 and H. angustatum

CBS 236.34

Fungal Diversity

Byssosphaeria schiedermayeriana MFLUCC10-0100

Byssosphaeria schiedermayeriana SMH 3157

Byssosphaeria schiedermayeriana GKM 152N

Byssosphaeria salebrosa SMH 2387

Byssosphaeria musae MFLUCC 11-0146

Byssosphaeria siamense MFLUCC10-0099

Herpotrichia diffusa CBS 250.62

Byssosphaeria jamaicana SMH 3085

Byssosphaeria jamaicana SMH 1403

Byssosphaeria rhodomphala GKM L153N

Byssosphaeria rhodomphala SMH 3086

Bertiella macrospora IL 5005

Byssosphaeria villosa GKM 204N

Pseudostrickeria muriformis MFLUCC13-0764

Pseudostrickeria ononidis MFLUCC14–0949

Phragmocephala atra MFLUCC15-0021

Phragmocephala garethjonesii MFLUCC15-0018

Sarimanas pseudofluviatile MAFF 239465

Sarimanas shirakamiense MAFF 242969

Sarimanas shirakamiense MAFF 244768

Aposphaeria corallinolutea CBS 131287

Aposphaeria corallinolutea CBS 131286

Herpotrichia macrotricha SMH 269

Herpotrichia macrotricha GKM 196N

Herpotrichia vaginatispora MFLUCC13-0865

Monotosporella tuberculata CBS 256.84

Herpotrichia juniperi CBS 200.31

Muriformistrickeria rubi MFLUCC15-0681

Melanomma rhododendri ANM 73

Melanomma pulvis-pyrius CBS 124080

Melanomma pulvis-pyrius CBS 109.77

Melanomma pulvis-pyrius CBS 371.75

Aposphaeria populina CBS 350.82

Aposphaeria populina CBS 543.70

Xenostigmina zilleri CBS 124108

Mycopappus aceris CBS 124109

Xenostigmina zilleri CBS 115685

Xenostigmina zilleri CBS 115686

Beverwykella pulmonaria MFLUCC15-0183

Beverwykella pulmonaria CBS 283.53

Pseudotrichia mutabilis SMH 5288

Pseudotrichia mutabilis SMH 1541

Prosthemium betulinum VM20040721

Pleomassaria siparia CBS 279.74

Herpotrichia parasitica CBS 451.73

Thyridaria rubronotata CBS 419.85

Massariosphaeria phaeospora CBS 611.86

Helicascus nypae BCC 36752

Morosphaeria ramunculicola JK 5304B

Trematosphaeria pertusa CBS 122368

Trematosphaeria pertusa CBS 122371

Bambusicola splendida MFLUCC 11-0439

Bambusicola massarinia MFLUCC 11-0389

Kalmusia scabrispora KT 2202

Bimuria novae-zelandiae CBS 107.79

Phaeodothis winteri CBS 182.58

Sporidesmiella fusiformis HKUCC 10831

Karstenula rhodostoma CBS 690.94

Paraphaeosphaeria michotii CBS 591.73

Montagnula opulenta CBS 168.34

Massarina cisti CBS 266.62

Massarina eburnea CBS 473.64

Byssothecium circinans CBS 675.92

Keissleriella cladophila CBS 104.55

Katumotoa bambusicola MAFF 239641

Lentithecium fluviatile CBS 122367

Lentithecium aquaticum CBS 123099

Sporormiella minima CBS 524.50

Roussoella hysterioides CBS 125434

Roussoella nitidula MFLUCC 11-0182

Roussoella nitidula MFLUCC 11-0634

Thysanolaenae thailandica MFLUCC 11-0155

Thysanolaenae rubriostiolata MFLUCC 11-0138

Thysanolaenae rubriostiolata MFLUCC 11-0137

Thysanolaenae rubriostiolata MFLUCC 11-0156

Pseudotrichia guatopoensis SMH 4535

Salsuginea ramicola KT 2597.1

Aigialus mangrovis BCC 33564

Aigialus grandis BCC 18419

Astrosphaeriella aggregata KT 767

Astrosphaeriella aggregata KT 984

Acrocordiopsis patilii BCC 28167

Acrocordiopsis patilii BCC 28166

Astrosphaeriella bakeriana CBS 115556

Astrosphaeriella stellata MFLUCC10-0555

Astrosphaeriella stellata MFLUCC10-0095

Caryosporella rhizophorae JK 5302A

Hysterium angustatum CBS 123334

Hysterium angustatum CBS 236.34

98/1.00

100/1.00

89/1.00

51/1.00

81/0.95

57/0.95

98/0.97

98/1.00

83/1.00

86/0.90

100/1.00

69/1.00

74/1.00

77/1.00

100/0.96

100/1.00

97/1.00

95/1.00

56/-

84/1.00

74/-

93/1.00

89/0.99

93/0.99

100/1.00

60/0.99

100/1.00

97/1.00

98/1.00

100/1.00

98/1.00

100/1.00

58/-

100/1.00

64/0.93

97/-

50/0.95

87/1.00

100/1.00

52/-

100/1.00

98/1.00

51/-

100/1.00

99/1.00

100/1.00

55/-

99/1.00

100/-

100/1.00

0.1

Melanommataceae

Pleomassariaceae

Morosphaeriaceae

Thyridariaceae

Trematosphaeriaceae

Bambusicolaceae

Didymosphaeriaceae

Massarinaceae

Lentitheciaceae

Roussoellaceae

Aigialaceae

Outgroup

ML/BYPP

Clade

Fungal Diversity

9. Ascospores with a mucilaginous sheath ..........................12

10. Ascospores muriform...............................Pseudostrickeria

10. Ascospores without longitudinal septa…....…….......…11

11. Ascospores ellipsoid to fusoid, 2 to multi-septate, con-

stricted at the main septum……………..……….Melanomma

11. Ascospores broadly to narrowly fusoid, 3-septate, easily

separating into two parts at the primary septum

......................................................................................Ohleria

12. Ascospores muriform..........................Muriformistrickeria

12. Ascospores without longitudinal septa…….....…......…13

13. Ascospores broadly fusiform with rounded ends,

hyaline……….......…..…..…..…..…..…..…...…..Sarimanas

13. Ascospores navicular to obovoid, hyaline to

brown……..............................................…Asymmetricospora

14. Pathogenic on host, produceing multicellular, epiphyllous,

splash-dispersed propagules……….....................Mycopappus

14. Saprobic on host, without propagules………..........…..15

15. Conidiophores branched.................................................16

15. Conidiophores unbranched.............................................18

16. Conidiophores branched, verruculose,

brown..................................................................Xenostigmina

16. Conidiophores branched, hyaline...................................17

17. Conidia hyaline, aseptate...............................Aposphaeria

17. Conidia brown, septate...................................Exosporiella

18. Conidia oblong to cylindrical, clavate to

obclavate..........................................................Nigrolentilocus

18. Conidia obovoid, ellipsoidal to subglobose………...…19

19. Conidia pyriform to obovoid, 1– 3-sep-

tate…………………………………........…Monotosporella

19. Conidia ellipsoidal to subglobose, 4– 6-sep-

tate………………………………...………Phragmocephala

Accepted genera of Melanommataceae

Aposphaeria

Berk., Outl. Brit. Fung. (London): 315 (1860)

FacesofFungi number: FoF 00756.

Saprobic in terrestrial habitats. Conidiomata pycnidial, su-

perficial, solitary, globose to subglobose, black, shiny, uniloc-

ular. Conidiomata wall thin, composed of one layer of medi-

um brown, thick-walled cells of textura angularis. Ostiole

single, circular, central, depressed. Conidiophores hyaline,

branched at the base, septate above, short, cylindrical, and

formed from the inner w all cells of the py cnidial wall.

Conidiogenous cells enteroblastic, phialidic, integrated or dis-

crete, determinate, hyaline, smooth, with apertures apical or

lateral just below septa; collarett e occasionally prominent,

with minute channel, with thickened periclinal wall. Conidia

hyaline, aseptate, thin-walled, eguttulate, cylindrical or ellip-

soidal, smo oth. My celium immersed, branched, septate,

brown (Sutton 1980).

Type species: Aposphaeria pulviscula (Sacc.) Sacc.

Notes: Aposphaeria was introduced by Saccardo (1880)

without description and illustration. The type species

Aposphaeria pulviscula which traditionally has been

identified as Phoma pulviscula is also poorly known. Based

on the molecular phylogentic analysis, some species are no

longer considered to be members of Aposph aeria,for

instance, Boerema et al. (1996) examined the holotype of

A. fusco-maculans and considered it to be A. pulviscula,which

fits the generic concept of Paraconiothyrium and de Gruyter

et al. (2012) subsequently confirmed the classification based

on mo lecular dat a. In addition, some other species in

Aposphaeria or Aposphaeria-like species have also been re-

ported in Massariosphaeria (Farr et al. 1989; Leuchtmann

1984;Kirketal.2001; Tanaka and Harada 2004), but these

records are unsubstantiated with molecular data. The polyphy-

letic relationships of Aposp haer ia, Massariosphaeria and

Melanomma were documented by Wang et al. (2007); they

suggested that more asexual morphs should be incorporated

in future studies. However, de Gruyter et al. (2012)showed

clearly that A. populina and A. corallinolutea are

melanommataceous and several species in Aposphaeria have

been published as the asexual morph of Melanomma.Inthe

present multigene phylogenetic analysis the asexual morph

strains of A. corallinolutea (CBS 131286 and CBS

131286) and

A. populina (CBS 543.70 and CBS

350.82) are included. Aposphaeria populina (CBS

543.70 and CBS 350.82) clustered together with

Melanomma pulvis-pyrius (CBS 371.75) with 81 % ML

BS and 0.95 BY PP (Fig. 1). Aposphaeria corallinolutea

(CBS 131286 and CBS 13 1286) form a single clade.

Sequence data for the type sp ecies of Apos phaeria

pulviscula is unavailable in GenBank, so we refrain from

synonymizing Apos phaeri a under Melanomma without

molecular data, thus, we retain Aposphaeria as a genus

in Melanommataceae, pending further study.

Aposphaeria pulviscula (Sacc.) Sacc., Michelia 2(no. 6): 4

(1880)

Synonymy:

≡Phoma pulviscula Sacc., Michelia 1(no. 2): 259 (1878)

≡Coniothyrium pulviscula (Sacc.) Kuntze, Revis. gen. pl.

(Leipzig) 3(2): 459 (1898)

FacesofFungi number: FoF 00757, Figs. 2 and 3

Saprobic on plants in terrestrial habitats. Mycelium im-

mersed, branched, septate, brown. Conidiomata (56−)84–

133μm high×(30−)54–100μm diam. (x̅=93×87 μm, n=10),

pycnidial, gregarious, superficial or semi-immersed, globose

to subglobose, black, ostiolate. Ostiole single, circular, central.

Pycnidial wall 9–13μm (x̅=11μm, n=10), comprising a sin-

gle layer of thick-walled, brown cells of textura angularis.

Conidiophores 1–4μm, hyaline, septate, cylindrical, formed

from the inner cells of the pycnidial base wall.

Conidiogenous cells enteroblastic, phialidic, hyaline, smooth.

Conidia 2.3–2.8×0.8–1.4μm (x̅=2.5×1.2μm, n=10), hya-

line, aseptate, thin-walled, eguttulate, cylindrical or ellipsoi-

dal, smooth-walled.

Fungal Diversity

Material examined: ITALY, Padova, in Salix udensis

Trautv. & C. Meyer (Salicaceae), January 1878, Saccardo

(IMI 202557, isotype,asex-isotypeofPhoma pulviscula).

Asymmetricospora J. Fröhl. & K.D. Hyde, Sydowia 50(2):

183 (1998)

FacesofFungi number: FoF 00758

Saprobic on palm fronds. Sexual morph: Ascomata

pseudostromatic, solitary or in small groups, immersed and

appearing as slightly black domes on the surface of substrate,

lenticular in vertical section, multi- or rarely uni-locular, indi-

vidual locule with a flat base. Ostiole single, central, opening

without tissue differentiation. Peridium carbonaceous, thicker

at sides and apex, composed of a few layers of pigmented cells

of textura angularis; base thinner, composed of irregularly-

shaped, hyaline cells arranged in a textura globulosa or

textura prismatica. Hamathecium of long, persistent, filiform,

branching, pseudo pa raph ys es, anasto mo sing betwee n and

above asci, within a gelatinous matrix. Asci 8-spored,

bitunicate, fissitunicate, clavate, with a short pedicel, apically

rounded with ocular chambers. Ascospor es biseriate, hyaline,

becoming pale brown when senescent, navicular to obovoid,

straight or curved, sometimes asymmetric, 1-septate, upper

cell larger with a rounded end, basal cell with a tapering

end, constricted at septum, usually with a spreading mucilag-

inous sheath, smooth-walled. Asexual morph:

Undetermined.

Fig. 2 Aposphaeria pulviscula

(isotype, from slides). a

Herbarium label of Aposphaeria

pulviscula. b–d Vertical sections

through peridium. f–g

Conidiogenous cells. Scale bars:

b–c=50μm, d–e=25μm,

f–g=5μm

Fungal Diversity

Type species: Asymmetricospora calamicola J. Fröhl. &

K.D. Hyde

Notes: Asymmetricospora was introduced to accommodate

the single species A. calamicola as a monotypic genus

(Fröhlich and Hyde 1998). The characteristics that distinguish

Asymmetricospora from other melanommataceous genera are

the absence of a subiculum, presence of short, dark setae,

around the papilla, and its asymmetrical ascospores

(Fröhlich and Hyde 1998). Morphologically

Asymmetricospora can be distinguished from

Astrosphaeriella (Fröhlich and Hyde 1998)as

Asymmetricospora are commonly multi-loculate with a

simple ostiole and without tissue differentiation, asco-

spores are asymmetric and ascomata normally coalesce,

whereas, Astrosphaeriella has one locule per ascoma

and w alls are t hick, amorphous and carbonaceous. In

this study, it is recommended that Asymmetricospora

should be maintained in Melanommataceae on account

of the trabeculate pseudoparaphyses (Lumbsch and

Huhndorf 2010; Zhang et al. 2012;Hydeetal.2013;

Wijayawardene et al. 2014), but it is acknowledged that

fresh collections, epitypification and molecular analysis

should be carried out to confirm the definite taxonomic

placement.

Asymmetricospora calamicola J. Fröh l. & K.D. Hyde,

Sydowia 50(2): 184 (1998)

FacesofFungi number: FoF 00759, Fig. 4

Saprobic on dead rattans of Calamus caryotoides. Sexual

morph: Ascomata pseudostromatic, 650–870(− 990)

μm×(500–)855–1300(−1500) μm (x̅=795×1100μm, n=10),

solitary or in small groups, immersed, subepidermal and

appearing as slightly protruding black domes on the substrate

surface, lenticular in vertical section, multi- or rarely unilocu-

lar, individual locules 320–450μ

m diam×170–285(−320) μm

high (x̅=227×385μm, n=10), with a flattened base. Ostiole

opening in the center, surrounded by a pale brown ring, with-

out tissue differentiation. Peridium 30–72(−96) μm wide, up-

per layer carbonaceous, cells melanised and occluded, becom-

ing pale brown textura angularis inwardly. Base thinner, com-

posed of irregularly-shaped, hyaline cells of textura globulosa

or textura prismatica. Hamathecium of long

pseudoparaphyses, up to 1.4–1.6(−2.5) μm wide, persistent,

filiform, branching and anastomosing between and above as-

ci, embedded in a gelatinous matrix. Asci 135–210μm×25–

35 μm (x̅ =172 ×33 μm, n =10), 8-spored, bitunicate,

fissitunicate, clavate, with short pedicel, 20–25μm,apically

rounded with ocular chambers. Ascospores (30–)35–55×10–

15μm (x̅=42×13μm , n=10), biseriate, hyaline, becoming

pale brown when senescent, navicular to obovoid, straight or

usually curved, asymmetric, 1-septate, upper cell larger with a

rounded end, basal cell tapering at the end, constricted at sep-

tum, usually with a muci lagin ou s sheath, smoo th-w alled .

Asexual morph: Undetermined.

Material examined: AUSTRALIA, North Queensland,

Palmerston, Palmerston National Park, on dead rattan of

Calamus caryotoides A.Cunn. ex Mart. (Arecaceae), 15

March 1994, J. Fröhlich (HKU(M) JF293, holotype).

Bertiella (Sacc.) Sacc. & P. Syd., in Saccardo, Syll. fung.

(Abellini) 14: 19 (1899) Basionym:

=Bertia subgen. Bertiella Sacc., Syll. fung. (Abellini) 1:

584 (1882)

FacesofFungi number: FoF 00760

Saprobic on woody hosts. Sexual morph: Ascomata gre-

garious sometimes confluent and in small groups, superficial,

subglobose, with a flattened base, apapillate, black, carbona-

ceous. Peridium composed of few layers of pigmented cells of

textura angularis, lower peridium thinner, composed of elon-

gated, hyaline cells of textura globulosa. Hamathecium of

long trabeculate pseudoparaphyses, numerous, persistent, fili-

form, branching and anastomosing between and above the

Fig. 3 Aposphaeria pulviscula

(re-drawn from Sutton 1980,

isotype). a Conidia. b

Conidiophores. c Vertical section

of a conidioma. Scale bars:

a=5μm, b=10μm, c=25μm

Fungal Diversity

asci. Asci 8-spored, bitunicate, cylindric-clavate, with short

pedicel, apically rounded with ocular chambers. Ascospores

biseriate, hyaline, becoming pale brown when senescent,

fusiform, straight or curved, 1-septate, with 2–3 globules, be-

coming 3-septate when mature, constricted at septum,

smooth-walled. Asexual morph: Undetermined.

Fig. 4 Asymmetricospora calamicola (holotype). a, b Specimens and

herbarium label of Asymmetricospora calamicola. c, d Appearance of

pseudostromata on the host surface. Note the central ostioles. f Vertic al

section through the tissue at the periphery of the pseudostroma. e, g

Vertical sections through peri dium. h Immature ascus. i–l Asci with

ascospores. m Trabeculate pseudoparaphyses. n–r Ascospores. Note the

mucilaginous sheath. Scale bars: c=500μm, d =200μm, f =100μm,

g=20μm, e, h–m=10μm, n–r=5μm

Fungal Diversity

Type species: Bertiella macrospora (Sacc.) Sacc. &

Traverso

Notes: Bertiella was raised to generic rank by Saccardo and

Sydow (1899), with Bertiella macrospora as the generic type.

Eriksson and Yue (1986) re-examined the type specimen of

B. macrospora andassignedittoMassarina (as

M. macrospora (Sacc.) O.E. Erikss. & J.Z. Yue). Bertiella

macrospora is characterised by superficial ascomata,

cylindro-clavate asci and hyaline, 1-septate ascospores that

may become 3-septate and pale brown when senescent. The

woody habitat in particular suggested Bertiella has similarities

with Lophiostoma based on morphology, and thus, Hyde et al.

(2002) assigned Bertia macrospora to Loph iostoma as

L. bertiellum Aptroot & K.D. Hyde. Mugambi and

Huhndorf (2009) provided molecular data from a single iso-

late of B. macrospora, which clustered with Byssosphaeria in

Melanommataceae. This species differs from Byssosphaeria

in having a roughened ascomatal surface that lacks hairs or a

subiculum. The main difference is that no cephalothecioid cell

arrangement is reported for this fungus, which is considered a

prominent feature for this genus (Mugambi and Huhndorf

2009). Bertiella is placed in Melanommataceae in contrast

to Teichosporaceae where it was previously placed

(Lumbsch and Huhndorf 2007). In the present multigene phy-

logenetic analyses, the strain of B. macrospora (IL 5005)

forms a distinct clade, close to Byssosphaeria in

Melanommataceae.

Bertiella macrospora (Sacc.) Sacc. & Traverso, Syll. fung.

(Abellini) 19: 147 (1910).

Basionym:

=Bertia macrospora Sacc., Michelia 1(no. 8): 452 (1882).

FacesofFungi number: FoF 00761, Fig. 5

Saprobic on woody host. Sexual morph: Ascomata gre-

garious sometimes confluent or in a small groups, semi-

immersed to superficial, subglobose with a flattened base,

apapillate, black, carbonaceous. Peridium 60–70μm thick,

partly consisting of strongly melanized isodiametric cells of

textura angularis in the outer parts, becoming paler and not

occuluded inwardly, base peridium thinner, composed of elon-

gated hyaline cells of textura angularis. Hamathecium of 1–

2μm wide, long, numerous, persistent, fil iform, trabeulate

pseudoparaphyses, branching and anastomosing between

and above the asci. Asci 130–150 ×22 μm, 8-spored,

bitunicate, cylindric-clavate, with short pedicel, apically

rounded, with an ocular chamber. Ascospores 37–43×8–

9μm biseriate, hyaline, becoming pale brown when senescent,

fusiform, straight or curved, 1-septate, becoming 3-septate

when mature, constricted at septum, with 2–3 globules,

smooth-walled. Asexual morph: Undetermined.

Material examined: ITALY, Treviso, Consiglio, on wood

of Fagus sylvatica L. (Fagaceae), P. A. Saccardo (UPS,

holotype of Bertia macrospora).

Bicrouania Kohlm. & Volkm.-Kohlm., Mycol. Res. 94(5):

685 (1990)

FacesofFungi number: FoF 00763

Saprobic on marine algae. Se xual morph: Ascomata

gregarious, i mmersed, erumpe nt to sup erficial, black,

globose to subglobose, coriaceous, ostiolate, papillate

or ep api lla te, periphysat e. Peridium thick at apex, thin-

ner at base, 2-layered. Hamathecium composed of long,

colourless, branching trabeculate pseudoparaphyses. Asci

8-spored, bitunicate, fissitunicate, cylindrical to clavate,

pedicellate, lacking an ocular chamber. Ascospores uni-

seriate or partially overlapping uni-seriate, reddish

brown to brown, ellipsoid with pointed to rounded ends,

1-septate, constricted at darkened septum, with lower

narrow and longer cell, smooth-walled. Asexual morph:

Undetermined.

Fig. 5 a, d–f Massarina

macrospora re-drawn from

Eriksson and Yue (1986). a

Ascomata on the surface of Fagus

sylvatica (Fagaceae). d Brown,

3-septate ascospores. e Hyaline,

1-septate ascospore. f Bitunacate

asci. b, c, g Bertiella macrospora

(IL 5005), re-drawn from

Mugambi and Huhndorf (2009).

b Vertical section of peridium. c

Pseudoparaphyses. g Bitunicate

asci. Scale bars: c=5μm, d,

e=10μm, f–g=20μm

Fungal Diversity

Type species: Bicrouania maritima (P. Crouan & H.

Crouan) Kohlm. & Volkm.-Kohlm.

Notes: Bicrouania was established to accommodate the

type species Bicrouania maritima which was previously iden-

tified as Didymosphaeria maritima (P. Crouan & H. Crouan)

Sacc. by Kohlmeyer and Volkmann-Kohlmeyer (1990). It is

characterized by superficial ascomata lacking a clypeus, thick-

walled asci and its association with marine algae (Kohlmeyer

and Volkmann-Kohlmeyer 1990). Bicrouania is presently

monotypic and placed in Melanommataceae (Lumbsch and

Huhndorf 2007, 2010; Jones et al. 2009; Zhang et al. 2012;

Hyde et al. 2013; Wijayawardene et al. 2014). However, this

genus lacks molecular data. Based on the melanommataceous

character and trabeculate pseudoparaphyses, we presently

maintain the genus in Melanommataceae.

Bicrouania maritima (P. Crouan & H. Crouan) Kohlm. &

Volkm.-Kohlm., Mycol. Res. 94(5): 685 (1990).

Synonymy:

≡Sphaeria maritima P. Cr ouan & H. Crouan, Florule

Finistère (Paris): 27 (1867)

≡Didymosphaeria maritima (P. Crouan & H. Cr ouan)

Sacc., Syll. fung. (Abellini) 1: 703 (1882)

≡Didymosphaerella maritima (P. Crouan & H. Crouan)

Cooke, in Fuckel, Grevillea 17(no. 82): 29 (1889)

FacesofFungi number: FoF 00764, Fig. 6

Saprobic on Halimione portulacoides L.. Sexual morph:

Ascomata (300–)320–440μm high×365–450(−460) μm

diam. (x̅ =385×425μm, n =10), gregarious, immersed,

erumpent to superficial, black, globose to subglobose, coria-

ceous, ostiolate, epapillate or papillate, periphysate. Peridium

38–47

μm thick laterally, up to 70μm thick at apex, thinner at

base, 2-layered, with outer wall composed of irregular pseu-

doparenchymatous, thick-walled, brown cells of textura

intricata and inner layer composed of delicate, large,

colourless cells of textura angularis. Hamathecium composed

of transversely septate, long, 1–1.5μm broad, colourless,

branching, trabeculate pseudoparaphyses, anastomosing be-

tween a nd above the asci. Asci 17 0–225(−240)×17.5–

22.5μm (x̅ =200×21μm, n =10), 8-spored, bitunicate,

fissitunicate, cylindrical to clavate, pedicellate, with a sco-

spores arranged in a cluster, without an ocular cham ber.

Ascospores 22–26×12–15μm (x̅=24×13μm, n=10), oblique-

ly uni-seriate or partially overlapping, reddish brown to

brown, ellipsoidal, with pointed to rounded ends, 1-septate,

thick-walled, slightly constricted at the darkened septum, low-

er cell narrow and longer, smooth-walled. Asexual morph:

Undetermined.

Material examined: FRANCE, Ael len Finistere, on

Halimione portulacoides (Amaranthaceae), Crouan and

Crouan, Herb. J. Kohlmeyer 3097 (IMI 330806, isotype of

Sphaeria maritima).

Byssosphaeria Cooke, Grevillea 7(no. 43): 84 (1879)

FacesofFungi number: FoF 00765

Saprobic on woody angiosperms. Sexual morph:

Ascomata solitary, scattered or sometimes gregarious, glo-

bose, subglobose or ovoid, spreading on su bstrate as a

subiculum, coriaceous, ostiolate, wall black, roughened.

Ostiole single, flattened, yellow, red or orange coloered at

pore. Peridium 2-layered, with outer layer composed of irreg-

ular, thick-walled, brown to dark brown cells of textura

epidermoidea, with inner layer composed of small thin-

walled light brown cells. Hamathecium thick with long, tra-

beculate pseudoparaphyses, embedded in mucilage, anasto-

mosing between and above asci. Asci 8-spored, bitunicate,

fissitunicate, thick-walled, cylindrical to clavate, with short

pedicel, apically rounded with an ocular chamber.

Ascospores uni-seriate or bi-seriate, pale brown, fusiform, ta-

pering at both ends, mostly straight, sometimes slightly

curved, 1-septate, slightly constricted at the septum, minutely

verrucose. Asexual morph: Pycnidia formed in culture.

Conidiogenous cells phialidic, lining cavity.Conidiahyaline,

ellipsoid or subglobose (Barr 1984).

Type species: Byssosphaeria keithi i (Berk. & Broome)

Cooke (Table 3)

Notes: Byssosphaeria was introduced by Cooke and

Plowright (1879), typified by Byssosphaeria keithii based on

its superficial ascomata seated on a Btomentose subiculum of

interwoven threads^, which includes various spec ies of

Sphaeria and Byssisedae. Byssosphaeria is a widespread ge-

nus, comprising approximately 42 species (Kirk et al. 2015),

with Pyrenochaeta or Chaetophoma-like asexual morphs

(Barr 1984; Hawksworth et al. 1995; Samuels and Müller

1978;Hydeetal.2011; Zhang et al. 2012). The type species

B. keithii was previously treated as a synonym of

B. schiedermayeriana (Sivanesan 1971), while

B. schiedermayeriana was synonymized as

H. schiedermayeriana Fuckel (von Arx and Müller 1975;

Bose 1961;Luttrell1973; Müller and von Arx 1962;

Sivanesan 1971). Barr (1984) found that

B. schiedermayeriana occurs exclusively in tropical regions

or greenhouse environments in temperate regions and

accepted a relatively narrow generic concept of

Herpotrichia,thus,Byssosphaeria were revived in

Melanommataceae.Barr(1990a) gave a detailed description

of Byssosphaeria which is characterized by superficial

ascomata with bright yellow, orange or red flat apices around

the ostiole, with dependan t hyphal appendages that merge

with the subiculum below and hy aline ascospores. Zhang

et al. (

2012) observed that the specimens examined in their

study had orange, greenish and flat apices. Byssosphaeria is

readily distinguishable from Herpotrichia on account of the

trabeculate pseudoparaphyses, as well as presence of a

subiculum (Hyde et al. 2000). This was confirmed by five loci

phylogenetic studies, which revealed this to be a monophylet-

ic genus clustering in Melanommataceae (Mugambi and

Huhndorf 2009; Zhang et al. 2012;Hydeetal.2013). The

Fungal Diversity

present study revealed that Byssosphaeria musae is close to

B. salebrosa (Cooke & Peck) M. E. Barr and

B. schiedermayeriana (Fuckel) M. E. Barr with 74 % ML

BSand1.00BYPP(Liuetal.2015). Our new species

Byssosphaeria siamense formed a clade allied with B. musae

in Melanommataceae (69 % ML BS and 1.00 BY PP). We

also describe and illustrate one fresh collection of

B. schiedermayeriana from Thailand. Multigene phylogenetic

analysis (Fig. 1) indicates that all Byssosphaeria species clus-

ter in Melanommataceae with strong support.

Byssosphaeria keithii (Berk. & Broome) Cooke [as

‘keitii’], Cooke and Plowright (1879)

Basionym:

=Sphaeria keithii Berk. & Broome Ann. Mag. nat. Hist.,

Ser. 4 17: 144 (1876)

FacesofFungi number: FoF 00766, Fig. 7

Saprobic on old rope. Sexual morph: Ascomata 350–

520(−600) μm high×(320−)400–620μm diam. (x̅=450×

550μm, n=10), scattered or sometimes gregarious, globose,

subglobose to turbinate or ovoid, spreading on substrate as

subiculum, coriaceous, ostiolate, black and shiny, wall black,

roughened. Ostiole flat, epapillate with a pore, with reddish

orange or greenish tinge at pore. Peridium 53–88μm thick (x̅=

72μm, n=10), 2-layered, outer layer composed of irregular,

thick-walled, brown to dark brown cells of textura

epidermoidea, cells up to 3μm diam., inner layer composed

of small, thin-walled, pale brown cells, up to 5μm diam.,

merging into pseudoparaphyses. Hamathecium of dense,

thick, long, trabeculate pseudoparaphyses, 1–2μm broad, em-

bedded in mucilage, anastomosing between and above asci.

Asci 80 –112(−150)×10–13μm (x̅=105×12μm, n=10), 8-

spored, bitunicate, fissitunicate, cylindro-clavate to clavate,

pedicel normally 15–20μm long, sometimes up to 65μm in

mature asci. Ascospores bi-seriate above and uni-seriate or

overlapping uni-seriate below, 25 – 32(− 40) μm ×5.5–

7.5(−10) μm, pale brown, fusoid, tapering at both ends, most-

ly straight, sometimes slightly curved, 1-septate, becoming 3-

septate after discharge, with hyaline appendages at each acute

to subacute end, in some mature spores without appendages,

slightly constricted at the septum, minutely v errucose.

Asexual morph: Pyrenochaeta sp. (Barr 1984; Samuels and

Müller 1978;Hydeetal.2011). Pycnidia 70–500 μmdiam.

Conidiogenous cells phialidic, lining cavity, 5–8×4–6μm to

5–10×3–6μm. Conidia 2.5–3.5(−4)×1.5–2(−

3) μm,hyaline,

ellipsoid or subglobose (Samuels and Müller 1978).

Material examined: IRELAND, Dublin, Glasnevin

Botanic Garden, on old rope, June 1872, W. Keith, ex herb.

Berkeley (K(M) 108784, holotype of Sphaeria keithii Berk.

&Broome).

Byssosphaeria siamensis Boonmee, Q. T ian & K.D. Hyde,

sp. nov.

Index Fungorum number: IF551430, Facesoffungi

number: FoF 01026; Figs. 8 and 9

Holotype:MFLU10–0099

Etymology: In re ference to the occurrence in Siam

(Thailand).

Saprobic on decaying wood. Sexual morph: Ascomata

501–692μm high×561–720μm diam., superficial, globose

to subglobose, solitary to gregarious, dark brown, setose, co-

riaceous, apically ostiolate. Ostiole single, central, with a

Table 3 Difference between Byssosphaeria species discussed in this study

Species Size Septation Ascospore colour Host References

Ascospores

(μm)

Asci (μm) Ascomata

(μm)

B. jamaicana (Sivan.)

M.E. Barr

25–35×7–880–120×12–15 340–500 1–3 Light to clear

brown

On decorticated rooting wood Barr 1984

B. keithii (Berk. &

Broome) Cooke

25–32×

5.5–7.5

80–112 ×10–13 350–620 1–3 Pale brown On old rope Berkeley and

Broome 1876

B. musae Phookamsak

& K.D. Hyde

30–33×5–6125–135×12–

430–630 1–3 Hyaline to

light brown

On leaf sheath of Musa sp.

(Musaceae)

Liu et al. 2015

B. rhodomphala

(Berk.) Cooke

18–23×

6–7.5

85–120×10–13 220–500 1–(3–) Light brown On wood and periderm

of various trees

Barr 1990a

B. salebrosa (Cooke &

Peck) M.E. Barr

40–50× 7–9120–150×

13–16.5

440–800 1–3(−5) Hyaline to

light brown

On dead roots of Acer spicatum

(Sapindaceae)

Barr 1984

B. schiedermayeriana

(Fuckel) M.E. Barr

32–42× 5–8100–150×

12–15

500–825 1–3(−5) Light brown On rotting logs and branches,

endocarps of coconut

Barr 1984

B. villosa (Samuels &

E. Müll.) Boise

30–40×

8–12

140–200×

17–25

340–470 1–2 Hyaline to brown On bark and decorticated wood

of angiospermous trees

Samuels and

Müller 1979



Fig. 6 Bicrouania maritima (isotype, slides). a, b Herbarium slides and

packet. c Vertical section through ascoma. d, f Vertical section through

peridium. e Vertical section through ascoma. Note the central ostiole. g

Immature ascus. h–k Asci with ascospores. Note the bitunicate asci. l

Pseudoparaphyses. m–q Ascospores. Scale bars: c=50μm, d–f=25μm,

g–k=10μm, l–q=5μm

Fungal Diversity

Fig. 7 Byssosphaeria keithii (holotype). a–d Specimens and herbarium

label. e–f Ascomata on the surface of substrate. g Vertical section through

ascoma. h Immature ascus. i, j Asci with ascospores. k Asci, note the

rounded apex with ocular chamber. l Hand-cut section through peridium.

m Hamathecium. n–o Ascospores. Scale bars: e=500μm, f=200μm,

l, g=50μm, h–k, m–o=5μm

Fungal Diversity

pore-like opening, orange to yellow around pore, epapillate.

Peridium 39–42μm wide, 3-layered, outer layer comprising

dark brown, thick irregular pseudoparenchymatous cells; mid-

dle texture with brown cells of textura angularis, and inner

layer composed of light brown cells of texura prismatica.

Hamathecium 1–2μm wide, comprising dense, trabeculate,

filiform, anastomosing, septate, pseudoparaphyses, embedded

in a gelationous matrix. Asci 112–148×10–16μm (x̅=129×

12.5μm, n=20), 8-spored, bitunicate, cylindrical clavate or

saccate, pedicellate, with a 14.5–24μm long pedicel, apically

rounded with ocular c hambers. Ascospores 40 .5–50×7–

11 μm(x̅=45×8.5μm, n=20), bi-seriate, hyaline to pale yel-

low, oblong fusiform, 1(−3)-septate, constricted at the septa,

slightly curved, surrounded by a gelatinous, fusiform sheath

and thickened at both ends, bearing delicate hyaline append-

age at ends, wall smooth or verrucose. Asexual morph:

Undetermined.

Culture characteristics: Ascospores germinating on MEA

within 12 h. Colonies on MEA, fast growing, reaching up to

10 mm diam. in 7 days at 28 °C. Mycelium superficial, ini-

tially white, later becoming light yellow, hairy, with entire

edge.

Material examined: THAILAND, Chiang Rai, Muang,

Ban Pu Kaeng Waterfall, on decaying wood of unidentified

host, 5 January 2010, S. Boonmee PK01 (MFLU 10–0029);

extype living culture MFLUCC 10–0099=IFRD 2176.

Fig. 8 Byssosph aeria siamensis a Decaying woody substrate. b

Ascomata formed on substrate. c Section through ascoma. d Peridium.

e Pseudoparaphyses. f–h Asci. i Close up of thickened apex. j Short

pedicel. k–m Ascospores. Scale bars: a =10 mm, b=500μm,

c=200μm, d=20μm, e, i– j=5μm, f– h=50μm, k–m =10μm

Fungal Diversity

Notes: This fungus was collected on decaying wood, has

ascomata which are superficial, surrounded by setae and

browntodarkbrown. Morphologically, Byssosphaeria

siamensis is similar to Byssosphaeria musae, but ascospores

of B. musae have a wing-like appendages near the central

septum. Multi-gene analyses shows that B. siamensis groups

with B. musae (69 % ML BS and 1.00 BY PP). Hence we

introduce a new species, B. siamensis, to accommodate our

collection (Fig. 1).

Byssosphaeria schiedermayeriana (Fuckel) M.E. Barr,

Mycotaxon 20(1): 34 (1984)

Basionym:

= Herpotrichia schiedermayeriana Fuckel, Jb. nassau. Ver.

Naturk. 27–28: 27 (1874) [1873–74]

Facesoffungi number: FoF 01027, Figs. 10 and 11

Saprobic on decaying wood. Sexual morph: Ascomata

423–511μm high×478–537μm diam., solitary to gregarious,

superficial, with basal subiculum, globose to subglobose, api-

cal ostiolate, covered by thick, branched hyphae, with rough-

ened, black wall. Ostiole single, central, epapillate, orange and

greenish around pore. Peridium 40–50μm wide, 3-layered,

outer layer composed of brown thick-walled cells of textura

epidermoidea, middle layer composed of large pale brown

cells of textura angularis, inner layer composed of small hy-

aline cells, merging into pseudoparaphyses. Hamathecium 1–

1.5(−2) μm wide, comprising dense, filiform, long trabeculate

pseudoparaphyses, anastomosing and branching between and

above asci. Asci 100–135×10–13μm (x̅=119.5×11μm

, n=

10), 8-spored, bitunicate, cylindrical-subclavate or saccate,

pedicel 30–35 μm long, apically thickened and rounded with

negligible ocular chamber. Ascospores 36–49×9–12μm (x̅=

46×10μm, n =10), bi-seriate, hyaline to pale brown,

cylindrical-fusiform, slightly curved, 1(−3)-septate, constrict-

ed at the septa, surrounded by a gelatinous sheath, with hya-

line appendages at each acute to subacute end or appendages

lacking, smooth-walled. Asexual morph: Coelomycetous

formed in culture. Pycnidia 70–500μm diam, in concentric

rings or in the aerial mycelium, globose, light brown to black,

surrounded by long hyphal-like hair, ostiolate. Pyenidial wall

of textura angularis. Conidiogenous cells 5–10×3–6μm,

phialidic, lining cavity, nearly globose or more elongated.

Conidia 2.5–3.5(−4)×1.5– 2( −3) μm, hyline, ellipsoid or

subglobose (Samuels and Müller 1978).

Cultural characteristics: Ascospores germinating o n

MEA within 12 h. Colonies on MEA, slow growing, reaching

5 mm diam. after 7 days at 28 °C. Mycelium superficial, pri-

mary white, later becomeing light yellow, hairy, with entire

edge.

Material examined: THAILAND, Chiang Mai, Mae

Taeng, Mae Sae Waterfall, on decaying wood of unidentified

tree, N 19°14′ E 98°39, 962 msl., 7 September 2009, S.

Boonmee MS-01 (MFLU 10–0030, reference speciesmen

designed here); living culture MFLUCC 10–0100=IFRD

2178.

Notes: This fungus was collected on decaying wood from

Thailand. Morphologically, it is similar to Byssosphaeria

schiedermayeriana, and multi-gene analysis shows that this

strain groups with B. schiedermayeriana

. Thus, we designate

a reference specimen (Ariyawansa et al. 2014b), here for

B. schiedermayeriana (basionym Herpotrichia

schiedermayeriana).

Calyptronectria Speg., Anal. Mus. nac. B. Aires, Ser. 3 12:

412 (1909)

FacesofFungi number: FoF 00767

Fig. 9 Cultures of

Byssosphaeria siamensis a

Germinating ascospore. b, c

Colonies on MEA. d Vegetative

hyphae in culture. Scale bars:

a=10μm, b, c=10 mm, d=5μm

Fungal Diversity

Saprobic on host in terrestrial habitats. Sexual morph:

Ascomata solitary, scattered, or sometimes in small groups,

immersed, erumpent to semi-immersed, lenticular to

subglobose, ostiolate. Ostiole small, with slightly protruding

papilla. Peridium 2-layered, outer layer composed of pseudo-

parenchymatous cells, thick walled, inner layer composed less

pigmented cells, merging with pseudoparaphyses.

Hamathecium of long, trabeculate pseudoparaphyses,

branching and anastomosing, embedded in a gelatinous

matrix. Asci 8-spored, sometimes 4-spor ed, bitunicate ,

fissitunicate, cylindrical to clavate, with a short, furcate pedi-

cel, apically rounded w ith an ocular chamber. Ascospores

broadly fusoid to fusoid with broadly to narrowly rounded

ends, hyaline, muriform with 3–5transversesepta,with1–3

longitudinal septa in the central cells when mature, slightly

constricted at the median septum, the upper cell often broader

than the lower one, smooth-walled. Asexual m orph:

Undetermined.

Fig. 10 Byssosphaeria schiedermayeriana (reference specimem) a Substrate with a close up of ascoma. b, c Section through ascoma and peridium. d

Pseudoparaphyses. e, f Asci. g–j Ascospores. Scale bars: a, b=200μm, c–f=20μm, g–j=10μm

Fungal Diversity

Notes: Calyptronectria was introduced by Spegazzini

(1909) to accommodate C. argentinensis Speg. and

C. platensis Speg., with C. platensis as the generic type. The

genus is characterized by immersed ascomata, filiform, tra-

beculate pseudoparaphys es and hyaline, murifo rm asco-

spores, as well as a peridium that turns reddish brown in

KOH (Rossman et al. 1999). Subsequently, Barr (1983)

placed Calyptronectria indica Dhaware and Calyptronectria

ohiensis in Calyptronectria based on the bitunicate asci (Barr

1983; Rossman et al. 1999). However, Rossman et al. (1999)

transferred C. ohiensis to Thyridium (as Thyridium ohiense

(Ellis & Everh.) Rossman & Samuels) because of its pseudo-

parenchymatous peridium. Calyptronectria is a poorly known

genus which currently contains three species, viz.

C. argentinensis, C. indica and C. platensis (Kirk et al.

2015). However, it has not been studied with molecular data

and there is no sequence data for any species in Genbank.

Fresh collections are required for epitypified, and epi-extype

living cultures should be obtained for future phylogenetic

studies. In the present study, Calyptr onectria is morphologi-

cally well-defined by trabeculate pseudoparaphyses, and thus,

retained in Melanommataceae.

Type species: Calyptronectria platensis Speg., Anal. Mus.

nac. B. Aires, Ser. 3 12: 412 (1909)

FacesofFungi number: FoF 01038, Fig. 12

For description see Zhang et al. (2012).

Exosporiella P. Karst., Finlands mögelsvampar,

(Hyphomycetes fennici): 160 (1892)

Synonymy:

≡Phanerocorynella Höhn., in Falck, Mykol. Untersuch.

Ber. 1(3): 351 (1923)

≡Anomalemma Sivan., Trans. Br. mycol. Soc. 81(2): 328

(1983)

FacesofFungi number: FoF 01024

Saprobic or fungicolous in terrestrial habitats. Sexual

morph: Ascomata gregarious on prosenchymatous stroma

with a intertwined hyphae, superficial, black, coriaceous or

carbonaceous, ostiolate. Ostiole single, circular to papillate,

central, depressed. Peridium a single layer, composed of

brown or heavily p igmented, thick-walled cells of textu ra

angularis or textura globosa. Hamathecium comprising

dense, filiform, hyaline, persistent pseudoparaphyses. Asci 8-

spored, bitunicate, clavate, with a short ped icel, apically

rounded with an ocular chamber. Ascospores uni-seriate, par-

tially overlapping bi-seriate, brown, short fusif orm to

Fig. 12 Calyptronectria platensis from LPS 1209, holotype (redrawn

from Zhang et al. 2012) a, b Ascospores. c Asci with

pseudoparaphyses. Scale bars: a–c=10μm

Fig. 11 Culture of

Byssosphaeria

schiedermayeriana a

Germinating ascospore. b, c

Colonies on MEA, upper surface

and lower surface. Scale bars:

a=10μm, b–c=10 mm

Fungal Diversity

narrowly fusoid, with narrowly rounded ends, 1-septate, rarely

2- to 3-septate, deeply constricted at the mid septum, smooth-

walled. Asexual morph: Coelomycetous. Intertwined hyphae

mainly superficial, around the pseudothecia , composed of

branched, brown to black hyphae with integrated

conidiogenous cells, distinction between conidiophores and

hyphae inconspicuous. Hyphae pale brown in mass, smooth.

Conidiophores reduced to conidiogenous cells.

Conidiogenous cells holoblastic, phialidic, percurrent, sub-

cylindrical to geniculate, sub-hyaline to pale brown, thin-

walled, smooth-walled, formed from the mass of hyphae

adpressed to the outer wall layer of the ascomata wall.

Conidia initially hyaline, pale yellowish brown to brown at

maturity, oblong to fusoid, mostly straight, infrequently slight-

ly curved, usually with 4 transverse septa, slightly constricted

at the septa, slightly rounded at apex, rounded to abruptly

attenuated towards a more or less conspicuous tapering base

with an appendage, smooth-walled.

Type species: Exosporiella epochnii (Berk. & Broome)

Sivan.

Notes: Berkeley and Broome (1866) described and illus-

trated Sphaeria epochnii from Britain. The hyphomycetous

Epochniella fungorum, which is the asexual morph was con-

firmed in that study and Epochniella fungorum has now been

transferred to Exosporiella. Sphaeria epochnii has previously

been transferred to many different genera. Saccardo (1878)

placed the sexual morph Anomalemma in Melanomma as

M. epochnii (Berk. & Broome) Sacc., Massee (1887)trans-

ferred it to Byssosphaeria as B. epochnii (Berk. & Broome)

Cooke, and Keissler (1922)transferedittoChaetosphaeria as

C. epochnii (Berk. & Broome) Keissl where it was illustrated

with unitunicate asci. Ellis (1883)placeditinLeptosphaeria,

although these is no evidence in the literature to support this.

A collection of S. epochnii from Dunwish Forest, Suffolk in

England, matched well with the description of S. epochnii.

(Sivanesan 1983). The asexual morph Exosporiella,whichis

reported to commonly occur on Corticium species in Europe

(Sivanesan 1983), has been described and illustrated by Ellis

(1971). There is no doubt that the asexual Exosporiella and

sexual morph are linked, however, Melanomma has

Aposphaeria or Pseudospiropes

asexual morphs, which differ

from Exosporiella (Sivanesan 1983). In addition, the

prosenchymatous stroma in E. epochnii also readily distin-

guishes it from Melanomma which has superficial gregarious

perithecial ascomata usually on a thin subiculum (Sivanesan

1983). Superfic ially, the gregarious prosenchymatous

ascomata in Anomalemma resembles those in Cucurbitaria.

However, this genus possesses dictyosporous asco spores

compared wi th the ascospores in Anomalemma which are

brown with one transverse septate and later becoming 3-sep-

tate, distinguishing it from other pleosporalean genera. In ad-

dition, the habitat of Anomalemma is fungicolous (Sivanesan

1983). Aposphaeria epochnii has similar characters with

gregarious ascomata on intertwined hyphae and brown

ascospores as in Byssosphaeria, while its fungicolous habitat

is the most obvious distinction. Anomalemma is presently

monotypic. According to the principle of prioty,

Wijayawardene et al. (2014) proposed that the oldest name

Exosporiella should be used for this genus. Thus,

Anomalemma should be regarded as a synonym of

Exosporiella. In this study, we did not examine the type of

Exosporiella, however we found this asexual morph when we

examine the isotype of Anomalemma epochnii. There has

been no phylogenetic analysis on this poorly studied genus.

There are no squences in GenBank of either the sexual or the

asexual states. The pseudoparaph yses are a typical of

Melanommataceae and fresh collections and sequence data

analysis should show the placement of this genus.

Exosporiella fungorum (Fr.) P. Karst., Finlands

mögelsvampar, (Hyphomycetes fennici): 161 (1892)

Synonymy:

≡Sphaeria epochnii Berk. & Broome, Ann. Mag. nat. Hist.,

Ser. 3 18: 128 (1866)

≡Melanomma epochnii (Berk. & Broome) Sacc., Michelia

1(no. 1): 34 (1878)

≡Byssosphaeria epochnii (Berk. & Broome) Cooke,

Grevillea 16(no. 78): 36 (1887)

≡Anomalemma epochnii (Berk. & Broome) Sivan., Trans.

Br. mycol. Soc. 81(2): 328 (1983)

FacesofFungi number: FoF 01025, Figs. 13 and 14

Saprobic on wood and bark. Sexual morph: Ascomata

320–450(−530) μm high×170–280(−350) μ

m diam. (x̅=

380×245 μm, n=10), on a prosen chymatous stroma with

intertwined hyphae, superficial, black, coriaceous or carbona-

ceous, ostiolate, roughened at the apex. Ostiole single, circular

to papilla, central, depressed. Peridium of a single wall layer,

(30− )44–67μm wide, compose d of brown or heavily

pigmented, thick-walled cells of textura angularis or textura

globosa, cells up to 8×13μm diam. Hamathecium of dense,

long cellular pseudoparaphyses, 2–3μm broad, septate, fili-

form, hyaline, persistent, branching above the asci. Asci 75–

108(− 120)×9–13μm (x̅ =92×11μm, n =10), 8-spored,

bitunicate, fissitunicate, cylindro-clavate to clavate, short bul-

bous to furcate pedicel, up to 6–15(−25) μm long, apically

round, with an ocular chamber. Ascospores 10–15×3–5.5μm

(x̅=13×4μm, n=10), uni-seriate to partially overlapping bi-

seriate, brown, short fusiform to narrowly fusoid with narrow-

ly rounded ends, 1-septate, rarely 2- to 3-septate, deeply con-

stricted at the central septum, smooth-walled. Asexual

morph: Hyphomycet ous. Colonies growing on bark of

Epochnium fungorum. Intertwined hyphae mainly superficial,

around the pseudothecia, composed of branched, brown to

black hyphae, with integrated conidiogenous cells, without

distinction between conidiophores and vegetative hyphae.

Hyphae

pale brown in mass, smooth-walled. Conidiophor es

reduced to conidiogenous cells. Conidiogenous cells 2–4×2–

Fungal Diversity

Fig. 13 Exosporiella fungorum (isotype of Anomalemma epochnii). a

Specimens and herbarium label of Anomalemma ep ochnii. b, c

Appearance of gregarious ascomata on the host surface. d

Pseudoparaphyses. e–h. Vertical sections through peridium. Note the

conidiogenous cell and conidia cluster on the adjacent stroma (arrowed

in e, f) i, j Conidiogenous cell. k, l Conidia. m, n Immature asci with

trabeculate pseudoparaphyses. o Asci with ascospores. p, q Ascospores.

Scale bars: b=500μm, c=200μm, e, f=25μm, g, h=10μm, g–j=10μm,

d, i–q=5μm

Fungal Diversity

5μm (x̅=3×3.5μm, n=10), holoblastic, percurrent, phialidic,

sub-cylindrical to geniculate, sub-hyaline to pale brown, thin-

walled, smooth, arising from mycelium at surface layer of

pseudothecial wall. Conidia 8–10×2–5.5μm (x̅=22×4μm,

n=10), oblong to fusoid, mostly straight, infrequently slightly

curved, usually with 4 transverse septa, slightly constricted at

septa, initially hyaline, pale yellowish brown to brown at ma-

turity, with central two cells dark brown and terminal cells

light brown, slightly rounded at apex, rounded to abruptly

attenuated towards a more or less conspicuous tapering base,

caudate, smooth-walled.

Material examined: UNITED KINGDOM, England,

Bath (near), Warleigh, on Epochnium fungorum Fr.

(Sclerotiniaceae) on bark, March 1866. (K (M) 188746,

isotype of Anomalemma epochnii, ex herb. C.E. Broome).

Herpotrichia Fuckel, Fungi rhenani exsic., suppl., fasc. 7

(nos 2101–2200): no. 2171 (1868)

FacesofFungi number: FoF 00770

Parasitic or saprobic on woody hosts in terrestrial habitats.

Sexual morph: Ascomata scattered to gregarious, immersed,

erumpent to nearly superficial, wall black, coriaceous, spread-

ing on substrate as a subiculum, ostiolate, roughened. Ostiole

single, cen tral, flattened, papillate. Peridium thick at both

sides and apex, basal wall thinner, 2-layered, outer layer com-

posed of pigmented, thick-walled cells of textura angularis,

inner layer comprising light pigmented, thin-walled cells of

textura prismatica. Ham athecium comprising dense, long ,

pseudoparaph yses, septate , branching and anastomosin g

above asci, embedded in a gelatinous matrix. Asci 8-spored,

bitunicate, fissitunicate, cylindrical to cylindro-clavate, with a

furcate pedicel, apically rounde d with an ocular chamber.

Ascospores bi-seriate to partially overlapping uni-seriate, hy-

aline, then becoming pale brown at maturity, fusoid, ellipsoid

or oblong with narrowly rounded ends, 1-septate, deeply con-

stricted at the septum, the upper cell often broader than the

lower on e, wall smooth or verruculose . Asexual morph:

Pyrenochaeta or Pyrenochaeta-like (Sivane san 1984).

Pycnidia solitary or gregarious, formed in culture from a sin-

gle ascospore isolate (Bose 1961), superficial, with or without

hairs, globose to pyriform or hemispherical, ostiolate. Ostiole

single, papillate with a long or short neck. Peridium wall com-

posed of

textura angularis. Conidiogenous cells simple, cy-

lindrical, branched, p hialidic. Conidia hyaline, globose to

ovoid or cylindrical, straight or curved, one-celled, with or

without guttules.

Type species: Herpotrichia herpotrichoides (Fuckel) P.F.

Cannon

Notes: Herpotrichia wasestablishedbyFuckel(1868)

basedontwospeciesHerpotrichia rhenana Fuckel and

Herpotrichia rubi Fuckel without assigning a type.

Subsequently, Bose (1961)designatedH. rhenana as the lec-

totype species. Müller and von Arx (1962) and Sivanesan

(1971) agreed with this proposal. Based on the priority of

principles of nomenclature (Cannon 1982), Holm (1979)in-

dicated that H. rubi should be assigned as the generic type as it

was validly published 2 years before H. rhenana. However,

Sphaeria herpotrichoides (synonymous with H. rubi)isthe

earliest name and priority should be given to

S. herpotrichoides. Finally, H. herpotrichoides was intro-

duced as the type of the genus (Holm 1979; Cannon 1982).

Herpotrichia is a morphologically and phylogenetically well-

studied genus (Barr 1984;Bose1961;MüllerandvonArx

1962;Pirozynski1972; Samuels and Müller 1978;

Sivanesan 1971, 1984; Mugambi and Huhndorf 2009;

Zhang et al. 2012). Currently, about 110 species are included

in this genus (Kirk et al. 2015). Herpotrichia is characterized

by erumpent to superficial ascomata, clavate to cylindrical, 4–

8-spored asci and hyaline to pale brown, 1-septate ascospores

(Sivanesan 1984), as well a Pyr enochaeta asexual morph with

or without setae on the surface of the pycnidia.

Morphologically, Herpotrichia and Melanomma share simi-

larities in the immersed, erumpent to superficial ascomata,

usually embedded on a subiculum, while in Herpotrichia as-

cospores tend to initially start hyaline and 1-septate, later be-

coming brown with more septa. Pseudotrichia differs from

Herpotrichia in its rather large ascomata, often with com-

pressed apices, while Herpotrichia produces smaller ascomata

usually on a subiculum (Bose 1961; Barr 1984). Herpotrichia

is very similar to Byssosphaeria but lacks the reddish-orange

or greenish tint near the ostiole, but has blunt papilla. In order

Fig. 14 Exosporiella fungorum (redrawn from Seifert et al. 2011)

Fungal Diversity

to obtain a natural concept for Herpotrichia, Zhang et al.

(2012) proposed to consider several criteria such as host, lo-

cation and habitat. In the present study, we introduce a new

species Herpotrichia vaginaspora, which clustered with other

Herpotrichia species. Herpotrichi a diffusa (CBS 250.62),

Herpotrichia juniper (CBS 200.31), Herpotrichia

macrotricha (SM H 269 a nd GK M 196N), Herpotrichia

parasitica (CBS 451.73) as well as the new taxon were in-

cluded and analyzed with four genes (LSU, SSU, RPB2 and

EF-1α). The resulting phylogeny reveals that Herpotrichia

species closer in different clades. Herpotrichia diffusa clus-

tered in Byssosphaeria,whileHerpotrichia juniper and

Herpotrichia macrotricha are close to Melanomma pulvis-

pyrius. However, H. parasitica (CBS 451.73) forms a single

clade close to Mor osphaeriaceae. Therefore, sequences data

for the type species H. herpotrichoides is needed to resolve the

phylogenetic position of this genus.

Herpotrichia herpotrichoides (Fuckel) P.F. Cannon, Trans.

Br. mycol. Soc. 79(2): 338 (1982)

Synonymy:

≡Herpotrichia rubi Fuckel, Jb. nassau. Ver. Naturk. 23–24:

146 (1870)[1869–70]

≡Sphaeria herpotrichoides Fuckel, Fungi rhenani exsic.,

fasc. 10: no. 952 (1864)

FacesofFungi number: FoF 00771, Fig. 15

Parasitic or saprobic on wood of Rubus idaeus L.. Sexual

morph: Ascomata 200–420(−450) μm high×230–400(−500)

μm diam ( x̅=365×348μm, n=10), scattered to gregarious,

immersed, erumpent to nearly superficial, coriaceous, spread-

ing on substrate as a subiculum, ostiolate, wall black, rough-

ened. Ostiole single, central, flattened, with a pore-like ostiole,

papillate, lacking periphyses. Peridium thick at both sides,

30–45μm wide and apex thicker, up to 57μm, basal wall thin-

ner, 2-layered, outer layer composed of pigmented, thick-

walled cells of textura angularis, inner layer comprising light

pigmented, thin-walled cells of textura prismatica.

Hamathecium comprising dense, 2–3 μm broad, long,

pseudoparaph yses, septate , branching and anastomosin g

above asci, embedded in a gelatinous matrix. Asci 95–145×

12–14(−16) μm (x̅=125×13μm, n=10), 8-spored, bitunicate,

fissitunicate, cylindrical to cylindro-clavate, with a furcate

pedicel, which is 18–40μm long, with an ocular chamber.

Ascospores 17–25×(5.5–)7–8.7μm (x̅=21×7.4μm, n=10),

bi-seriate to partially overlapping uni-seriate, fusoid, ellipsoid

or oblong with narrowly rounded ends, hyaline, then becom-

ing pale brown when mature, 1-septate, deeply constricted

at the septum, the upper cell often broader than the

lower one, verruculose. Asexual morph: Pyrenochaeta

rhenana Sacc. (Sivanesan 1984). Pycnidia formed in

culture from ascospore isolate, globose to p yriform,

hairy, up to 200μm broad, with a short ostiolar apex.

Conidiophores 6–10×1 .5– 2 μm short, simple. Conidia

1.5–2.5×1–2μm, hyaline, 1-celled.

Material examined: GERMANY, in the Oestreicher

meadow forest, on Rubus idaeus L. (Rosaceae), very rarely

in the s pring, 1894, Fuckel (S-F208018, holotype of

Herpotrichia rubi).

Herpotrichia vaginatispora Q.Tian, Wanasinghe,

Camporesi & K.D. Hyde, sp. nov.

Index Fungorum number: IF551595 , FacesofFungi

number: FoF 01028, Fig. 16

Holotype: MFLU 15–1408

Etymology: vaginae (sheaths) + spora (spore) referring to

ascospore sheath.

Saprobic on dead stem of Trifolium sp. Sexual morph:

Asco mata 145–190μm high, 150–210μm diam. (x̅=178×

190μm, n =10), solitary, scattered, globose or flattened,

semi-immersed, dark brown to black, coriaceous, ostiolate,

smooth-walled. Ostiole single, central, papillate, without

periphyses. Peridium 12–21μm (x̅=18.5μm, n=7), compris-

ing 2 strata, outer strata composed of heavily pigmented,

thick-walled cells of textura angularis, inner strata comprising

light brown, thin-walled cells of textura angularis.

Hamathecium of dense, 1.5–2.1μm broad, long, septate,

pseudoparaphyses, branching and anastomosing above the as-

ci, embedded in a gelatinous matrix. Asci

85–96×14–16μm

(x̅=89.7×15.2μm, n=10), 8-spored, bitunicate, fissitunicate,

cylindrical to cylindric-clavate, bulbous pedicellate, apically

rounded with an ocular chamber. Ascospores 26.5–31.5×6.5–

7.5μm (x̅=29.4×7.2μm, n=10), uni-seriate, partly overlap-

ping or bi-seriate, hyaline, fusoid, ellipsoid or oblong with

narrowly rounded ends, 1-septate, deeply constricted at the

septum, the upper cell often broader than the lower one,

guttulate, smooth-walled, with a structured mucilaginous

sheath. Asexual morph: Undetermined.

Culture characteristics: Colonies on PDA reaching 15–

20 mm diam at 25 °C after 21 d, surface white in centre, pale

dirty white to buff in outer region, spreading with sparse aerial

mycelium and even, smooth margins; iron-grey below.

Material examined: ITALY, province of Arezzo, near

Badia Prataglia, on dead stem of Tri f ol i um sp.

(Leguminosae), 29 August 2013, Erio Camporesi IT 1436

(MFLU 15–1408, holotype ; HKAS 88736, isotype), ex-

type living culture MFLUCC 13–0865=KUMCC.

Notes: Herpotrichia vaginatispora is introduced based on

morphological charaters and phylogenetic analysis.

Morphologic ally, Her potrichia vaginatispora is similar to

H. macrotricha in having immersed to erumpent ascomata

with a papillate ostiole, cylindrical to cylindric-clavate asci

and hyaline ascospores. Ho we v er, t he as co sp o re s i n

H. vaginatis por a are guttulate, usually with a mucilagi-

nous sheath, which is distinct from other Herpotrichia

species. In the present phylogenetic analysis,

H. vaginatispora clustered close to H. macrotricha

(GKM 196N, SMH 269) and forms a distinct clade in

Melanommataceae (Fig. 1).

Fungal Diversity

Mamillisphaeria K.D. Hyde et al., Nova Hedwigia 62(3–

4): 514 (1996)

FacesofFungi number: FoF 00772

Saprobic on woody host in freshwater environments.

Sexual morph: Ascomata scattered, sometimes gregarious,

superficial, conical, carbonaceous, ostiolate. Ostiole single,

Fig. 15 Herpotrichia herpotrichoides (holotype of Herpotrichia rubi)

a, b Specimens and herbarium label. c–e Appearance of ascomata on the

host surface. e Vertical section through ascoma. g–i Vertical section of

peridium which comprises cells of textura angularis. j Immatured ascus.

k– l Asci with ascospores. m Trabeculate pseudoparaphyses. n–o

Ascospores. Scale bars: g=50μm, h, i=25μm, j–l=10μm, m–o=5μm

Fungal Diversity

central, papillate. Peridium thick, comprising several layers of

compressed, densely packed, thin-walled, hyaline cells with

hyaline, palisade-like cells occuring at the periphery.

Hamathecium comprising long, branched or simple, septate,

trabeculate pseudoparaphyses, embedded in a gelatinous

matrix. Two types of asci and ascospores exist in the same

ascoma: TYPE 1: Asci 8-spored, cylindro-clavate, bitunicate,

fissitunicate, short-pedicellate, apically rounded with an ocu-

lar chamber. Ascospores 2–4-seriate, hyaline, ellipsoidal, sep-

tate, constricted at septum, with pad-like, m ucilaginous

Fig. 16 Herpotrichia

vaginatispora (holotype). a, b

Appearance of scattered ascomata

on the host surface. c Ve rt ical

section through ascoma. d

Vertical section through peridium.

Note the asexual projections. e

Pseudoparaphyses. f–i Asci with

ascospores. j–m Ascospores, note

the mucilaginous sheath when

stained with India ink. n

Germinating ascospore. Scale

bars: a=500μm, b=200μm,

c=50μm, d, n=20μm,

f–m=10μm, e=5μm

Fungal Diversity

appendage at each end and with some mucilage associated

around the spore. TYPE 2: Asci 8-spored, cylindrical,

bitunicate, fissitunicate, pedicellate, with an ocular chamber

and faint apical ring. Ascospores 1–2-seriate, brown, ellipsoi-

dal-fusoid, with a thin mucilaginous sheath. Asexual morph:

Undetermined.

Type species: Mamillisphaeria dimorphospora K.D. Hyde

et al.

Notes: Mamillisphaeria was introduced by Hyde et al.

(1996b) as a monotypic genus based on its bitunicate,

fissitunacate asci, trabeculate pseudoparaphyses and dimor-

phic ascospores to accommodate a freshwater species named

M. dimorphospora from tropical Australia. Both types of as-

cospores have mucilaginous sheaths (Hyde et al. 1996b). The

different function between two kin d of ascospores in

M. di morphospora was dis cussed by Hyde and Wong

(1999). The hyaline ascospores, which germinate within

72 h on PDA, may be adapted for attachment, enhanced col-

lision and provide nutrient for germination (Hyde et al. 1996b;

Zhang et al. 2012), The brown ascospores did not germinate

even after 7 days and may help withstand desiccation and UV

radiation during aerial dispersal (Hyde et al. 1996a, b;Zhang

et al. 2012). According to examination of M. dimorphospora,

the hyaline ascospors are easy to find in each fruiting body,

whereas the brown ascospores appear to be present in some

specimens only. Mami llisphaeria dimorphospora was re-

ferred to Massariaceae previously by Barr (1990a), while

Hyde and Wong (1999) temporarily assigned this species to

Melanommataceae on account of the trabeculate

pseudoparaphyses embedded in mucilage. Currently, in the

absence of molecular data but considering the distinctive mor-

phological characteristics, Mamillisohaeria is retained in

Melanommataceae.

Mamillisphaeria dimorphospora K.D. Hyde et al., Nova

Hedwigia 62(3–4): 515 (1996)

FacesofFungi number: FoF 00773, Fig. 17

Saprobic on submerged wood in a stream. Sexual morph:

Ascomata 450–650(−700) μm high×950–1350μm diam. (x̅=

585×1100μm, n=10), scattered or sometimes gregarious, in

small groups, superficial, conical, carbonaceous, ostiolate, un-

der pseudostroma that forms a thin layer on the host surface.

Ostiole single, central, papillate. Peridium 10–25μm thick,

comprising several layers of compressed, densely packed,

thin-walled, hyaline cells, with hyaline, palisade-like cells,

occuring at the periphery. Hamathecium of dense, septate,

trabeculate pseudoparaphyses, up to 1μm broad, hyaline,

branching and anastomosing above the asci, embedded in a

gelatinous matrix. Two types of asci and ascospores exist in

the same ascoma: TYPE 1: Asci 185–300×40–90μm (x̅=

255×72μm, n=10), 8-spored, cylindro-clavate, bitunicate,

fissitunicate, short-pedicellate, apically rounded with an ocu-

lar chamber (13μm wide×5μm high). Ascospores 65–85×

20–49 μm(x̅=76×35μm, n=10), 2-4-seriate, hyaline, light

brown when mature, ellipsoidal, 1-septate, constricted at the

central septum, with pad-like, mucilaginous appendage at

each end and with some mucilage associated around the spore;

TYPE 2: Asci 155–240×8–15μm (x̅=185×12μm, n=10), 8-

spored, cylindrical, bitunicate, fissitunicate, pedicellate, with

an ocular chamber and faint apical ring. Ascospores 30–42×

5–9μm (

x̅=35×7μm, n=10), 1-2-seriate, brown, dark brown

when mature, ellipsoidal-fusoid, 1-septate, constricted at the

septum, guttulate, smooth-walled, surrounded by a thin muci-

laginous sheath. Asexual morph: Undetermined.

Material examined: AUSTRALIA, Queensland, Cow

Bay, 16°13′ S, 145°26′ E, on submerged wood in a stream,

April 1995, Tamsin Hyde and K.D. Hyde (BRIP 22967a,

isotype)

Melanomma Nitschke ex Fuckel, Jb. nassau. Ver. Naturk.

23–24: 159 (1870) [1869–70]

Synonymy:

≡Moriolopis Norman ex Keissl., (1927)

FacesofFungi number: FoF 00774

Saprobic or parasitic on wood and bark in terrestrial hab-

itats. Sexual morph: Ascomata gregarious, i mmersed,

erumpent to nearly superficial, globose to subglobose, broadly

or narrowly conical, often laterally flattened, black, coria-

ceous, ostiolate. Ostiole single, central, papillate, often some-

what puckered or sulcate. Peridium comprising of two layers,

outer layer co mpri sing, s mall, hea vily pig ment ed, thi ck-

walled cells of textura angularis and inner layer comprising

lightly pigmented to hyali ne, thin-walled cells of textura

angularis. Hamathecium of dense, thickened, filamentous,

septate, trabeculate pseudoparaphyses, branching and rarely

anastomosing above the asci. Asci 8-spored, bitunicate,

fissitunicate, cylindrical to fusoid, with a short pedicel, apical-

ly rounded with an ocular chamber. Ascospores uniseriate and

partially overlapping, pale brown to brown, broadly fusoid to

fusoid, rounded at both ends, straight or slightly curved, 3-

septate, constricted at the septa, the second cell from the apex

slightly wider than the others, smooth-walle d. Asexual

morph: Coelomycetous. Conidiomata pycnidial, superficial,

globose, black, ostiolate, circular, central. Mycelium im-

mersed, branched, septate, brown. Pycnidial wall comprising

two layers, outer thick-walled brown cells of textura angularis

and inner layer comprising thin-walled hyaline cell of irregu-

lar issue. Conidiophores hyaline, septate, cylindrical, formed

from the inner cells at base of the pycnidial wall.

Conidiogenous cells enteroblastic, phialidic, hyaline, smooth.

Conidia hyaline, aseptate, thin-walled, eguttulate, cylindrical

or ellipsoidal, smooth-walled (Sutton 1980).

Type species: Melanomma pulvis-pyrius (Pers.) Fuckel

Notes: Melanomma was established by Fuckel (1870)and

is typified by Melanomma pulvis-pyrius

Winter (1887)based

on its small, carbonaceous ascomata, having hyaline or

brown, 2–3-septate ascospores (Chesters 1938; Fuckel 1870)

and is the familial type of Melanommataceae. Saccardo

Fungal Diversity

(1883) supplemented the description and synonymized many

species of Sphaeria as Melanomma. This was followed and

confirmed by Barr (1990a, b), and Melanomma pulvis-pyrius

was selected as the lectotype species (Barr 1990a; Chesters

Fungal Diversity

1938). In addition, a detailed circumscription of Melanomma

was given by Barr (1990a), with 20 species (Kirk et al. 2001),

while presently 299 epithets are listed in Kirk et al. (2015)of

which 62 are considered to belong to other genera (Speices

Fungorum 2015). The placement of the genus has been clar-

ified based on molecular and morphological data. Anamorphs

reported for this genus are Aposphaeria, Nigrolentilocus,

Phoma-like and Pseudospiropes (Chesters 1938; Sivanesan

1984;Hydeetal.2011). Aposphaeria agminalis Sacc. was

demonstrated as the asexual morph of M. pulvis-pyrius, which

is characterized by depressed to raised, cottony to woolly,

grey, colonies with rhizoidal margins in culture (Zhang et al.

2012). One of the diagnostic characters of

Melanommataceae is the trabeculate pseudoparaphyses

(Sivanesan 1984;Barr1990a; Liew et al. 2000;Kirk

et al. 2001). This genus has been well-studied phyloge-

netically. There are nearly 50 hits for sequence data in

GenBank. Zhang et al. (2008) observed the neotype of

M. pulvis-pyrius, and also collected new specimens of

M. pulvis-pyrius. After comparing morphological charac-

ters, they epitypified the species and provided an ex-

epitype culture (IFRDCC 2044). The only difference

between the neotype and epitype specimens is in the

peridium, which is thicker in the neotype (Zhang et al.

2008). Phylogenetic analysis based on sequences of four

genes (LSU, SSU, RPB2 and EF-1α) indicated that

Melanomma pulvis-pyrius clustered in a well-supported

clade with Byssosphaeria, Herpotrichia and

Pleomassaria and represented a separate f amily (or fam-

ilies c ompri sing Melanommataceae)(Zhangetal.2008;

Mugambi and Huhndorf 2009;Zhangetal.2012; Hyde

et al. 2013; Wijiyawardene et al. 2014). In this study,

M. pulvis-pyrius (CBS 124080, CBS 109.77 and CBS

371.75), M. rhododendri (ANM 73), Aposphaeria

populina (CBS 543.70 and CBS 350.82) and

A. corallinolutea (CBS 131286 and CBS 131287) were

retrieved from GenBank and analyzed with four loci.

Multigene analyses (Fig. 1) indicate that asexual morph

Aposphaeria populina is related t o M. pulvis-pyrius,as

theyclusteredtogetherwith81%MLBSand0.95BY

PP.However,sinceAposphaeria corallinolutea formed a

single cl ade close to Herpotrichia,itisproposedthat

Aposphaeria corallinolutea is not the asexual morph of

Melanomma. It is not possible to synonymize

Aposphaeria under Melanomma until the molecul ar data

of the type species of Melanomma and Aposphaeria is

obtained, thus Aposphaeria and Melanomma are

retained separately as sister genera in

Melanommataceae.

Melanomma pulvis-pyrius (Pers.) Fuckel, Jb. nassau. Ver.

Naturk. 23–24: 160 (1870) [1869–70]

Synonymy:

≡ Sphaeria pulvis-pyrius Pers., Syn. meth. fung.

(Göttingen) 1: 86 (1801).

Faces of Fungi number: FoF 00775, Fig. 18

Saprobic on wood and bark of Salix capre in terrestrial

habitats. Sexual mor ph: Ascomata 210–450(− 500) μm

high×(200–)250–450μm diam. (x̅=328×352μm, n=10),gre-

garious, immersed, erumpent to nearly superficial, globose to

subglobose, broadly or narrowly conical, often laterally flat-

tened, black, coriaceous, roughened and irregular, ostiolate.

Ostiole single, central, papillate, often somewhat puckered

or sulcate, lacking perip hyses. Peridium 65–85μm wide,

thicker at the base, up to 200μm,comprisingtwolayers,outer

layer composed of pigmented, thick-walled cells of textura

angularis, inner layer thinner, 5–8 μm

, composed of

irregularly-shaped lightly pigmented to hyaline, thin-walled

cells of textura angularis. Hamathecium comprising 1–2μm

broad, dense, thickened, septate, trabeculate

pseudoparaphyses, branching, rarely anastomosing the asci.

Asci 90–135×6.5–8(−9) μm (x̅ =117×7.5μm, n =10), 8-

spored, bitunicate, fissitunicate, cylindrical to fusoid, with a

short, bulbous to furcate pedicel, to 20μm long , apica ll y

rounded with an ocular chamber. Ascospores 14–17(−19)×

4.5–6μm (x̅=17×6μm, n=10), obliquely uni-seriate and over-

lapping uni-seriate, pale brown to brown, broadly fusoid to

fusoid, rounded at both ends, straight or slightly curved, 3-

septate, slightly constricted at the septa, with the second cell

from apex slightly wider than others, smooth-walled. Asexual

morph: Coelomycetous. Pycnidia 200–300μm in diam,

scattered or crowded, immersed to superficial, globose-coni-

cal, black, carbonaceous, shortly papillate. Pycnidial wall

consisting of 4–6 layers of dark brown cells, outer thick-

walled cells enclosing a pseudoparenchymatous layer of hya-

line cells. Conidiogenous cells flask-shaped, with apex of

young conidiogenous cells rounded . Conidia 2–3.5×1.5–

2.5μm, hyaline, ellipsoid or somewhat oblong, rounded at

the ends, guttulate.

Material examined: FRANCE, Ariège, Rimont, Saurine,

on bark of Salix caprea L. (Salicaceae), 10 April 2008,

Jacques Fournier (IFRD 2001, epitype).

Monotosporella S. Hughes, Can. J. Bot. 36: 786 (1958)

FacesofFungi number: FoF 00782

Colonies effuse, scatter ed, black. Mycelium immersed.

Conidiophores arranged in groups or arise singly from the



Fig. 17 Mamillisphaeria dimorphospora (isotype). a Specimens and

herbarium label. b, c Ascomata on host surface. Note the small papilla.

d Vertical hand-cut section of ascoma. e Vertical hand-cut section through

peridium. f Trabeculate pseudoparaphyses in a gelatinous matrix. g–j

Asci with ascospores (TYPE 2), note the cylindrical asci. k Both types

of asci in centrum. l Ascus with ascospores (TYPE 1), note the hyaline,

ellipsoidal ascospores with pad-like mucilaginous appendage at each end.

m– n Ascospores (TYPE 2), note the brown, ellipsoidal-fusoid

ascospores. o–p Ascospores (TYPE 1), note the pale, ellipsoidal-fusoid

ascospores. Scale bars: b=1000μm, c=500μm, d=100μm, e=50μm, f–

l=20μm, m–p=10μm

Fungal Diversity

substrate, erect or arcuate, red-dish brown to black, smooth-

walled, unbranched, 1– 2-septate. Conidiogenous cells

monoblastic and form pyriform conidia which arise singly at

the apex. Conidia solitary, pyriform to obovoid, 1–3septate,

usually two-septate, not constricted at the septa, dark brown to

black with hyaline cells at the base, smooth-walled;

Type species: Monotosporella setosa (Berk. & M.A.

Curtis) S. Hughes, Can. J. Bot. 36: 787 (1958), Fig. 19

Fig. 18 Melanomma pulvis-pyrius (epitype). a, b Appearance of

gregarious ascoma ta on the h ost surface. c Vertical section through

ascomata. d, f Vertical sections through peridium. e Filamentou s,

branching, septate pseudoparaphyses. g–j Ascospores. k–m Asci with

ascospores. Note the bitunicate asci with an ocular chamber. Scale bars:

a=500μm, b=200μm, c=100μm, f=50μm, d=20μm, e, g–m=10μm

Fungal Diversity

Synonymy:

≡Monotospora setosa Berk. & M.A. Curtis, in Berkeley,

Grevillea 3(no. 27): 110 (1875)

≡Phragmocephala setosa (Berk. & M.A. Curtis) E.W.

Mason & S. Hughes, Naturalist, London: 97 (1951)

≡Monotosporella setosa var. macrospora S. Hughes, N.Z.

Jl Bot. 16: 338 (1978)

Notes: Monotosporella was introduced by S. Hughes

(1958) and is typified by M. setosa Berk. & M. A. Curtis.

Monotosporella has a worldwide distribution and occurs on

decaying wood in freshwater and on terrestrial decaying

woody plants (Rao a nd de Hoog 1986;Rajaetal.2007;

Sivichai et al. 1998;Tsuietal.2001). Ellis (1959)synony-

mized Monotosporella under Brachysporiella, a genus which

has conidiopho res which branch at th e apex. Ho wever,

Monotosporella is characterized by unbranched conidio-

phores (Matsushima 1975). Subsequently, Hughes (1958)in-

dicated that Monotosporella should be distinct from

Brachysporiella, since lateral branches have not been de-

scribed in any of the published Monotosporella species.

Monotosporella was listed in Melanommataceae by

Wijayawardene et al. (2012). However, Seifert et al. (2011)

hadstatedthatMonotosporella is probably heterogenous.

Presently, 12 species epithets are listed in Kirk et al. (2015)

of which eight are considered to belong to Monotosporella

(Speices Fungorum 2015). The molecular data of

M. tuberculata (CBS 256.84) places it in Melanommataceae

(Schoch et al. 2009; Zhang et al. 2009b; Zhang et al. 2012;

Hyde et al. 2013; Wijiyawardene et al. 2014). In the present

study, the strain (CBS 256.84) is included and it is close to

Herpotrichia in Melanommataeae. However, it is still neces-

sary to recollect the type species in order to fully establish the

phylogenetic position of genus Monotosporella.

Muriformistrickeria Q. Tian, Wanasinghe, Camporesi &

K.D. Hyde, gen. nov.

Index Fungorum number: IF551596 , FacesofFungi

number: FoF 01035

Etymology: muriformis (muriform) + strickeria,referring

to muriform ascospores.

Type species: Muriformistrickeria rubi Q. Tian,

Wanasinghe, Camporesi & K.D. Hyde, sp. nov.

Saprobic on dead herbaceous branches in terrestrial habi-

tats. Sexual morph: Ascomata solitary, scattered, sometimes

gregarious, flattened, semi-immersed to erumpent, dark

brown to black, coriaceous, smooth, ostiolate. Ostiole single,

central, pore-like, falttened. Peridium 2-layered, outer layer

composed of irregular, thick-walled, brown to black cells of

textura angularis; inner layer comprising oblong, thin-walled,

hyaline to pale brown cells of textura prismatica.

Hamathecium la cking pseudopar aphyses. Asci 8-spored,

bitunicate, fissitunicate, cylindrical, short pedicellate, apically

rounded, with an ocular chamber. Ascospores uni-seriate or

overlapping, ellipsoidal, muriform, 4–6transverselyseptate,

with 2–4 vertical septa, initially hyaline, becoming brown at

maturity, conical and narrowly rounded at the ends, yellow to

brown, constricted at the center, lower cell narrow and longer,

smooth-walled, with a mucilaginous sheath. Asexual morph:

Conidiomata pycnidial, gregarious, superficial, black, cir-

cular. Mycelium immersed, branched, septate, brown.

Pycnidial wall comprising two layers, outer thick-

walled brown cells of textura angularis and inner l ayer

comprising thin-walled hyaline cell of irregular issue.

Conidiophores hyaline, septate, cylindrical, f ormed from

the inner cells of the pycnidial base wall.

Conidioge nous ce lls enteroblastic, phialidic, hyaline,

smooth. Conidia hyaline, aseptate, thin-walled,

eguttulate, cylindrical or ellipsoidal, smooth-walled.

Notes: A new monotypic genus Muriformistrickeria is in-

troduced here to accommodate Muriformistrickeria rubi.The

muriform ascospore is the diagnostic character to distinguish

Muriformistrickeria from other genera in Melanommataceae,

but it is m orpholog ically sim ilar wi th Pseudostrickeria.

Muriformistrickeria has muriform ascospores constricted at

the median transverse septa, while conidia in

Pseudostrickeria species are not constricted. Based on phylo-

genetic analyses, M. rubi (MFLUCC 15–0681) is close to

Herpotrichia group. Compared with the morphological char-

acters, Herpotrichia has hyaline, fusoid, 1-septate ascospores.

Thus, with no other genus to accommodate this taxon, our

collections are introduced as a new genus and species.

Fig. 19 Monotosporella setosa (Conidiophores and conidia redrawn

from Crous 1978). a, b, c, e from DAOM 96020; a, f from DAOM

109262; d from DAOM 93803

Fungal Diversity

Muriformistrickeria rubi Q. Tian, Wanasinghe,

Camporesi & K.D. Hyde, sp. nov.

Index Fungorum number: IF551597 , FacesofFungi

number: FoF 01036, Fig. 20

Etymology: Named after the host Rubusi sp..

Holotype:MFLU15–1409

Saprobic on dead herbaceous branches. Sexual morph:

Asco mata 200–230μm high, 240–265μm diam. (x̅=210×

247μm, n=10), solitary, scattered, or sometimes gregarious,

globose or flattened, semi-immersed to erumpent, dark brown

to black, coriaceous, smooth, ostiolate. Peridium 21–32μm

(x̅=28.5μm, n=7), 1-layered, composed of irregular, thick-

walled, brown to black cells of textura angularis.

Hamathecium lacking pseudoparaphyses. Asci 85–130×18–

24 μm (x̅ =110×22μm, n =10), 8-spored, bitunicate,

fissitunicate, cylindrical to cylindric-clavate, short pedicellate,

Fig. 20 Muriformistri ckeria rubi (holo type) a, b Appearance of

gregarious ascoma ta on the h ost surface. c Vertical section through

ascoma. d Vert ical section through peridium. e – g Verti cal sec tions

through conidioma. h– k Asci with ascospores. l–m Conidiogenous

cells. n– k As cospores, note the mucilaginous sheath staining

with India ink. r Conidia. Scale bars: c =100μm, d, f–g=20μm,

e=50μm, h–k=10μm, l– r=5μm

Fungal Diversity

apically rounded, with an ocular chamber. Ascospores 19–

27×9–13μm (x̅=25×11μm, n=10), uni-seriate or overlap-

ping uniseriate, sometimes bi-seriate, mostly ellipsoidal,

muriform, 4–6transverselyseptate,with2–4verticalsepta,

initially light yellow, becoming yellowish-brown at maturity,

conical and narrowly rounded at the ends, constricted at the

center, lower cell narrow and longer, smooth-walled, with a

thick mucilaginous sheath. Asexual morph: Conidiomata (56

− )87–146μm high×(30− )300– 360μm diam. (x̅ =121×

347μm, n=10), pycnidial, gregarious, superficial or semi-im-

mersed, black, flattened, circular, in the central. Mycelium

immersed, branched, septate, brown. Pycnidial wall 17–

27μm (x̅=24μm, n=10), comprising two layers, outer layer

of thick-walled brown cells of textura angularis and inner

layer comprising thin-walled irregula r, hyaline cells.

Conidiophores 1–4μm, hyaline, septate, cylindrical, formed

from the inner cells of the pycnidial base. Conidiogenous cells

enteroblastic, phialidic, hyaline, smooth. Conidia 3.3–4×2–

2.4μm (x̅=3.7×2.2μm, n=10), hyaline, aseptate, thin-walled,

eguttulate, cylindrical or ellipsoidal, smooth-walled.

Culture characteristics: Colonies on PDA reaching 20–

25 mm diam at 25 °C after 21 d, surface olivaceous-grey,

spreading with moderate aerial mycelium, reverse iron-grey.

Material examined: ITALY, Province of Arezzo [AR],

Montemezzano-Stia, on dead branch of Rubus sp.

(Rosaceae) , 23 September 2013, E. Campor esi IT 1460

(MFLU 15–1409, holotype, HKAS–88737, isotype), ex-

type living culture, MFLUCC 15–0681, KUMCC15-0079.

Mycopappus Redhead & G.P. White, Can. J. Bot. 63(8):

1430 (1985)

FacesofFungi number: FoF 00776

Pathogenic with leaf spots variable in shape, subcircular to

irregular, with pale green or dark brown margin. Propagules

conspicuous, on leaf spots or near the spots, numerous,

scattered to aggregated, white to pale brown, erumpent from

the cuticle, easily abscised, composed of stromatic base and

long or short filamentous appendages. Stromatic base dichot-

omously branched. Long-type appendage (filamentous hy-

phae) cylindrical, with obtuse apex, 2–8-septate. Short-type

appendage (claviform hyphae) cylindrical to clavate, aseptate

or 1–3(−4)-septate. Both appendage types forming together

along with stromatic base.

Type species: Mycopappus alni (Dearn. & Barthol.)

Redhead & G.P. White

Notes: Mycopappus was introduced by Redhead and White

(1985) for asexual species producing multicellular,

epiphyllous , spla sh-disp ersed propagule s previ ously de -

scribed as Cer cosporella alni Dearness & Bartholomew and

C. aceris Dearness & Batholomew. Presently, there are four

species in this genus (Kirk et al. 2015), viz. M. aceris (Dearn.

& Barthol.) Redhead & G.P. White, M. aesculi C.Z. Wei, Y.

Harada & Katum., M. alni (Dearn. & Barthol.) Redhead &

G.P. White, and M. quercus Y. Suto & M. Kawai. All species

are leaf pathogens (Redhead and White 1985). Crous et al.

(2009) re-examined Xenostigmina and showed that

Mycopappus may be a synanamorph and allied to Seifertia

in Pleosporales. Furthermore, Crous et al. (2013) revealed

that in their phylogenetic analysis Xenostigmina and

Mycopappus clustered in Phaeosphaeriaceae. However, in

the phylogenetic analysis of Phookamsak et al. (

2014),

Mycopappus clustered with Xenostigmina in a single clade

close to Melanommataceae and thus was excl uded fr om

Phaeosphaeriaceae. In the present study, the placement of

M. aceris (CBS 124109) was determined based on LSU,

SSU, RPB2 and EF-1α sequence data analysis. It demonstrat-

ed that M. aceris (CBS 124109) aligns with Xenostigmina

zilleri (CBS 124108, CBS 115685, CBS 115686) in a well-

supported clade (93 % ML BS and 0.99 BY PP). These two

genera are indeed synanamorphs, thus Mycopappus is retained

in Melanommataceae.

Mycopappus alni (Dearn. & Barthol.) Redhead & G.P.

White, Can. J. Bot. 63(8): 1430 (1985)

Synonymy:

≡Cer cosporella alni Dearn. & Barthol., Mycologia 9:362,

1917

FacesofFungi number: FoF 01029, Fig. 21

Leaf spots variable in shape. Propagules conspicuous, on

leaf spots or near the spots, numerous, scattered to aggregated,

white to pale brown, conical when filaments agglutinated, or

mop like in form when they spread, erumpent from the cuticle,

easily abscised. Propagules composed of stromatic base and

long or short filamentous appendages. Stromatic base dichot-

omously branched, interwoven cells, giving rise to numerous

long or short filamentous hyphae. Long-type appendage (fil-

amentous hyphae) cylindrical, with obtuse apex, 87.5–280×

9.5–14.5μm (x̅=230.5×12.5μm, n =20), 2–8-septate. Short-

type appendage (claviform hyphae) cylindrical to clava te,

55.0–92.5×8.0–13.5μm (x̅=87.3×10.5μm, n =20), aseptate

or 1–3(−4)-septate. Both appendage types intermingling, the

claviform hyphae could be immature filamentous hyphae.

Material examined: USA, Washington, Bremerton, on

leaves of Alnus rubra Bong. (Betu laceae), 9 September

1912, E. Bartholomew 4924 (IMI 8617, holotype

Navicella Fabre, Annls Sci. Nat., Bot., sér. 6 9: 96 (1879)

[1878]

FacesofFungi number: FoF 00777

Saprobic on dead wood in terrestrial habitats. Sexual

morph: Ascomata solitary or scattered, immersed to

erumpent, through the outer layer of the host tissue to nearly

superficial, coriaceous, black, subglobose and usually flat-

tened. Ostiole usually widely porate, with well-developed

neck, ostiolar canal filled with a tissue of hyaline cells.

Peridium 2 -layered, comprising strongly pigmented, dark

brown to black, thick-walled cells of textura angularis.

Hamathecium of dense, long, trabeculate pseudoparaphyses,

branching, anastomosing between and above the asci,

Fungal Diversity

embedded in mucilage. Asci bitunicate, fissitunicate, clavate

to sub-cylindrical, long pedicellate, apically rounded, with a

minute ocular chamber. Ascospores biseriate, partially over-

lapping, brown to chestnut-brown, ellipsoidal, with broadly

rounded ends, 5–8-septate, constricted at the septum, verru-

cose, with hyaline appendage at each end, smooth-walled.

Asexual morph: Undetermined.

Type species: Navicella julii Fabre

Notes: Navicella was introduced by Fabre (1879), and was

treated by Barr (1990a) who described the genus as character-

ized by immersed to erumpent ascomata, clavate or cylindrical

asci, trabeculate pseudoparaphyses and distoseptate asco-

spores. The genus Navicella is saprobic on bark, and consid-

ered to be closely related to Lophiostomataceae (Holm and

Holm 1988;Thambugalaetal.2015). However, Navicella is

distinguished from other genera in Lophiostomataceae on the

basis of its ascospores, which have conspicuously thickened

(distosepta) septa and lenticular lumina (Eriksson 1981;

Thambugala et al. 2015). Barr (1990a) placed it in

Massariaceae based on wide endotunica, thin apical ring

and distoseptate ascospores. However, this was not accepted

by Voglmayr and Jaklitsch (2011) and Zhang et al. (2012),

who indicated that the morphological characters of Navicella

did not match Massa riaceae. Navicella shares similarities

with Melanommataceae in having superficial g lobose to

subglobose, coriaceous ascomata with long, trabeculate

pseudoparaphyses and brown ascospores, b ut differs

from other genera of Melanommataceae in having ascospores

with euseptate primary septa, while in others septa are

distoseptate, with hyaline appendages at each end. Currently,

there are 116 Navicella species listed in Ind ex Fungorum

(Kirk et al. 2015), but no molecular data is available for phy-

logenetic analysis of any species. Based on morphological

characters, the taxonomy of Ariyawansa et a l. (2014a)is

followed to assign Navicella to Melanommataceae pending

phylogenetic investigation.

Fig. 21 Mycopappus alni

(holotype) a–c Specimens and

herbarium label. d Appearance of

gregarious propagules on host

surface. e–g Propagule composed

of stromatic base and long or

short filamentous appendages. h

Claviform hyphae i Filamentous

hyphae. Scale bars: e–i=20μm

Fungal Diversity

Navicella julii Fabre, Annls Sci. Nat., Bot., sér. 6 9: 96

(1879) [1878]

FacesofFungi number: FoF 01030, Fig. 22

Saprobic on dead wood in terrestrial habitats. Sexual

morph: Ascomata 750–890×460–600μm (x̅=810×520μm,

n=10), solitary or scattered, erumpent to nearly superficial,

black, subglobose and usually flattened, coriaceous, smooth-

walled. Ostiole widely porate, with well-develope d neck,

ostiolar canal filled with a tissue of hyaline cells. Peridium

33–45μm wide (x̅ =38μm, n=10), 2-layered, outer layer

comprising highly pigmented, dark brown to black, thick-

walled cells of textura angularis, inner layer comprising pale

brown to hyaline, thin-walled cells of textura angula ris.

Hamathecium of dense, 1–1.5μm (x̅=1.2μm, n=20) wide,

long, trabeculate pseudoparaphyses, branching and anasto-

mosing between and above the asci, embedded in mucilage

matrix. Asci 150–200×15–20μm (x̅=160×17μm, n=20), 8-

spored, bitunicate, fissitunicate, clavate to sub-cylindrical,

long pedicellate and apically rounded and thickened with an

ocular chamber. Ascospor es 55–75×8–12μm (x̅

=60×10μm,

Fig. 22 Navicella julii

(holotype). a, b Specimens and

herbarium label. c, d Appearance

of ascomata on the surface of

host. e Vertical section through

ascoma. f Hand-cut section

through peridium. g, h Asci with

ascospores. i Immature ascus. j

Trabeculate pseudoparaphyses.

k–n Ascospores. Scale bars:

e=100μm, f, j=10μm,

g–i=15μm, k–n=5μm

Fungal Diversity

n=40), biseriate, partially overlapping, brown to chestnut-

brown, ellipsoidal, 5–8-sept ate, primary septum euseptate,

and others distoseptate, verrucose, with hyaline appendage

at both ends. Asexual morph: Undetermined.

Material examined: FRANCE, Vaucluse, on dead wood,

H. Fabre (S-F71928, holotype).

Nigrolentilocus R.F. Castañeda & Heredia, Cryptog.

Mycol. 22(1): 13 (2001)

FacesofFungi number: FoF 00778

Hyphomycetous fungi on woody plants in terrestrial

habatats. Sexual morph: Melanomma-like sp.. Asexual

morph: Colonie s effuse, hairy, oliveceous, b rown to

dark brown o r black. Mycelium superficial, brown.

Hyphae branched, pale brown or brown, septate, smooth

or verrucose. Conidiophores conspicuous,

mononematous, solitary or fasciculate, septate, erect,

straight or flexuous, smooth or verrucose, cylindrical,

dark brown to black. Conidiogenous cells holoblastic

or polyblastic, pale brown to brown. Conidiogenous loci

large, thickened, protuberant and black, lenticular.

Conidia oblong to cylindrical, navicular, clavate to

obclavate, broadly fusiform to turbinate, always broadly

truncate and black at the base, euseptate,

acropleurogenous, pale brown to dark brown, wall

smooth or verrucose.

Type species: Nigrolentilocus africanus (B. Sutton) R.F.

Castañeda & Heredia

Notes: Nigrolentilocus was introduced by Castañeda et al.

(2001) after a revision of the asexual genus Pseudospiropes.A

sexual morph was noted but never described in either treat-

ments of the type species. This asexual genus accommodates

the species with discoid, broad, protuberant, black

conidiogenous loci and euseptate conidia. Nigrolentilocus

africanus, N. novae-zealandiae, N. variabilis, N. longipilus

and N. lotorus were transferred to Nigrolentilocus

(Castañeda et al. 2001). Recently, Nigrolentilocus amazonicus

J.S. Monteiro, Gusmão & R. F. Castañeda was introduced by

Monteiro et al. (2014) and currently, there are six species listed

in Kirk et al. (2015). Molecular data is lacking in GenBank

(2015). However, despite this, when Wijayawardene et al.

(2012) liste d the a sexual m orphs o f Melanommataceae,

Nigrolentilocus was included and subsequently reported by

Hyde et al. (2013) and Wijayawardene et al. (2014

). In the

present study, the holotype of Pseudospiropes africanus was

re-examined, and the sexual morph was observed on the host

surface, but no asexual morph was detected. The morpholog-

ical characters of the sexual morph do not fit the generic de-

scription of Melanomma, such as 1-septate ascospores and

long pedicles on the asci. The placement of Nigrolentilocus

cannot be con firmed, but based on the e vidence of the

previous authors, Nigrolentilocus is accepted in

Melanommataceae. Epitypification of the type species

N. africanus is required to verify the phylogenetic placement.

Nigrolentilocus africanus (B. Sutton) R.F. Castañeda &

Heredia, in Castañeda Ruiz et al., Cryptog. Mycol. 22(1): 15

(2001)

Basionym:

=Pseudospiropes africanus B. Sutton, Mycol. Pap. 167:

52 (1993)

FacesofFungi number: FoF 01031, Figs. 23 and 24

Hyphomycetous on stem o f Lobelia sp. in terrestrial

habatats. Asexual morph: Colonies effuse, hairy, olivaceous,

brown to dark brown or black. Mycelium superficial, brown,

immersed. Hyphae branched, pale brown or brown, septate,

smooth or verrucose. Conidiophores conspicuous,

mononematous, solitary or fasciculate, septate, erect, straight

or flexuous, smooth or verrucose, cylindrical, dark brown to

black. Conidiogenous cells holoblastic or polyblastic, pale

brown to brown. Conidiogenous loci enlarged, thickened, pro-

tuberant and black, lenticular. Conidia oblong to cylindrical,

navicular, clavate to obclavate, broadly fusiform to turbinate,

always broadly tr unc ate and black at the base, euseptate,

acropleurogenous, pale brown to dark brown. smooth or ver-

rucose-walled. Sexual morph: Ascomata 490–675(−780) μm

high×(300–)420–560μm diam. (x̅=589×515μm, n=10),gre-

garious, semi-immersed to erumpent, globose to subglobose,

often late rally flatte ned, black, coriaceous , roughened ,

ostiolate. Ostiole single, central, papillate, lacking periphyses.

Peridium 34–62μm wide, thinner at apex, 38–45

μm wide,

thicker at the base, up to 112μm, comprising two layers, outer

layer composed of pigmented thick-walled cells of textura

angularis, inner laye r thinner composed of irreg ularly-

shaped lightly pigmented to hyaline thin-walled cells of

textura angularis. Hamathecium comprising 2–3μm broad,

dense, thick, septate, pseudoparaphyses, branching, rarely

anastomosing above the asci. Asci 95–115× 10–11(−13) μm

(x̅=102×10.9μm, n=10), 8-spored, bitunicate, fissitunicate,

cylindrical to fusoid, with a long pedicel, apically rounded,

with an ocular chamber. Ascospores 13–16(−18)×4.5–6.5μm

(x̅=15.5×5.8μm, n=10), obliquely uni-seriate and bi-seriate,

light brown to brown, broadly fusoid to fusoid, both ends

rounded, straight or slightly curved, 1–3-septate, slightly con-

stricted at the septa, with second cell from the apex slightly

wider than others, smooth-walled.

Material examined: AFRICA, Tropical Southern Africa,

Malawi, Mt Mulanje, Chambe hut track, south-east on track to

Lichenya, on stem of Lobelia sp. (Campanulaceae), 13 April

1991, B.C. Sutton (MM42) (IMI 347112a, holotype of

Pseudospiropes africanus B. Sutton 1993).

Ohleria Fuckel, Fungi rhenani exsic., suppl., fasc. 7(nos

2101–2200): no. 2173 (1868), FacesofFungi number: FoF

00779

Saprobic on decaying roots in terrestrial habitats. Colonies

solitary or confluent of variable shape and size, black. Sexual

morph: Ascomata solitary, scattered or gregarious, superfi-

cial, carbonaceous, broadly or narrowly conical, with a

Fungal Diversity

flattened base, ostiolate. Ostiole single, central, papillate.

Peridium thicker at the apex, 2-layered, composed of

pigmented thick-walled cells of textura angularis,becoming

thinner towards the inner layers. Hamathecium comprising

long, branched or simple, septate, pseudoparaphyses, in a

slimy matrix encasing the asci. Asci 8 -spor ed, bitunicate,

fissitunicate, cylindrical, with short broad pedicel, apically

rounded with an ocular chamber. Ascospores bi-seriate, brown

to reddish brown, broadly fusoid to fusoid with broadly to

narrowly rounded ends, 3-septate, constricted at the median

septum and easily breaking into two conical partspores at the

primary septum, not constricted at the secondary septum,

smooth-walled. Asexual morph: Monodictys sp. from culture

of Ohleria brasiliensis. Conidiophores free, micronematous,

integrated, terminal. Conidiogenous cells holoblastic, cylin-

drical, determinate or less frequently percurrent. Conidia sol-

itary, brown, aerogenous, simple, muriform (Samuels 1980;

Hyde et al. 2011).

Type species: Ohleria modesta Fuckel

Notes: Ohleria was introduced by Fuckel (1868) and later

described and illustrated by Samuels (1980). Distinguishing

morphological characteristics for this genus are the

subglobose to conical ascomata, and phragmosporous asco-

spores tha t break into two parts at the median septum

(Samuels 1980). Clements and Shear (1931)treatedOhleria

as a synonym of Ohleriella,despiteOhleriella being a cop-

rophilous fungus. When t he ascomata and habitats are

Fig. 24 Nigrolentilocus africanus (conidia re-drawn from Castañeda

Ruiz et al. 2001)

Fig. 23 Nigrolentilocus

africanus (holotype of

Pseudospiropes africanus). a

Herbarium label and specimens.

b, c Appearance of gregarious

ascomata on the host surface. d, e

Vertical section through ascoma. f

Vertical sections through

peridium. g–i Asci with

ascospores. j Asci, note apically

rounded with an ocular chamber.

k Trabeculate pseudoparaphyses.

l–o Ascospores. Scale bars: d,

e=50μm, f=20μm, g–j=10μm,

k–o=5μm

Fungal Diversity

considered, Ohleria seems to be more closely related to

Melanomma and Trematosphaeria (Samuels 1980).

However, the differences in ascospores characters (bi-seriate,

easily breaking into two conical partspores in Ohleria vs

uniseriate ascospores not breaking in Melanomma) demon-

strate the distincion between Melanomma and Ohleria.Inad-

dition, Ohleria has been considered closely related to

Sporormia (Rehm 1912), and O. aemulans Rehm has been

transferred to Ohleria. According to Kirk et al. (2015), cur-

rently, there are 14 epithets included in this genus, while

Species Fungorum (2015) considers 13 species of Ohleria.

Some species of Ohleria are widespread, such as

O. brasiliensis reported from New Zealand, Brazil as well as

United States (Samuels 1980). Presently, there are no se-

quence data in GenBank. Therefore fresh collections for

DNA isolation, sequencing and phylogenetic analysis are nec-

essary to determine the natural classification of Ohleria.Asa

consequence, it is suggested that Ohleria should be retained in

Melanommataceae pending phylogenetic analysis.

Ohleria modesta Fuckel, Fungi rhenani exsic., suppl., fasc.

7(nos 2101–2200): no. 2173 (1868)

FacesofFungi number: FoF 00780, Fig. 25

Saprobic on the terrestrial host. Colonies growing on

decorticated decaying roots of Fagus sylvatica, solitary, con-

fluent of variable shape and size, black . Sexual morph:

Ascomata (200–)220–370μm high×(240–)285–400μm diam.

(x̅=342×385μm, n=10), solitary, scattered and gregarious in

small groups, erumpent to nearly superficial, black, carbona-

ceous, broadly or narrowly conical, with a flattene d base,

ostiolate. Ostiole single, central, with a conical protruding or

pore-like papilla. Peridium thicker at the apex, 26–45(

−55)

μm thick at both sides, 1-layered, composed of pigmented,

thick-walled cells of textura angularis. Hamathecium 1–

2μm broad, comprising long, dense, septat e, cellular

pseudoparaphyses, branching and anastomosing above the as-

ci. Asci 90–120×(5–)7–10μm (x̅=116×8.5μm, n=10), 8-

spored, bitunicate, fissitunicate, cylindrical, with short broad

pedicel, apically rounded, with an ocular chamber. Ascospores

12–20(−25)×3–5μm (x̅=18.7×4μm, n=10), bi-seriate and

uniseriate at the base, broadly fusoid to fusoid with broadly

to narrowly rounded ends, brown to reddish brown, 3-septate,

constricted at the median septum and easily breaking into two

conical partspores at the primary septum, constricted at the

middle septum, smooth-walled. Asexual morph:

Undetermined.

Material examined: GERMANY, on decorticated,

decaying roots of Fagus sylvatica L. (Fagaceae), collected

in autumn (G: F. rh. 2173, isotype).

Phragmocephala E.W. Mason & S. Hughes, Naturalist

(Hull), ser. 3, 1951: 97 (1951) FacesofFungi number: FoF

00783

Sexual morph: Undetermined. Asexual morph: Colonies

superficial, scattered, dark brown to black. Hyphae hyaline,

septate, smooth. Conidiophores macronematous, septate, un-

branched or branched, erect, dark brown at the base, pale

brown at apex, 4–8-septate. Conidiogenous cells holoblastic,

terminal, integrated, light brown to pale brown. Conidia 4–6

septate, ellipsoidal to subglobose, dark brown, pale brown at

apical and basal cells, sometimes often released with part of

conidiogenous cell.

Type species:

Phragmocephala elliptica (Berk. & Broome)

S. Hughes, N.Z. Jl Bot. 17(2): 164 (1979)

Notes: Phragmocephala was established by E.W. Mason &

S. Hughes (1951 ) and is typif ied by P. elliptica (Berk. &

Broome) S. Hughes w hich was originally described as

Monotospora elliptica from dead herbaceous hosts (Mason

and Hughes 1951; Whitton et al. 2012). Subsequently, Ellis

(1959, 1971) synonymized Phragmocephala in

Endophragmia due to the similarities in conidiogenesis.

However, Hughes (1979) reassigned the genus transferring

four species from Endophragmia. Presently, 14 species

epithets are listed in Index Fungorum (Kirk et al. 2015),

which a re considered to represent 9 s pecies (Species

Fungorum 2015). Su et al. (2015) described a new spe-

cies, P. garethjonesii (MFLUCC 15–0021) and presented a

new collection of P. atra (MFLUCC 15–0018) providing

molecular data. Phragmocephala was demonstrated to be a

distinct clade. Therefore, the taxonomy of Su et al. (2015)is

followed and the placement of Phragmocephala is deter-

mined to be in the family Melannomataceae.

Pseudostrickeria Q. Tian, Wanasinghe, Camporesi & K.D.

Hyde, gen. nov.

Index Fungorum number: IF551598 , FacesofFungi

number: FoF 01032

Etymology: In reference to the morphological similarity to

Strickeria Körb.

Type species: Pseudostrickeria muriformis Wanasinghe, Q.

Tian, Camporesi & K.D. Hyde, sp. nov.

Saprobic on dead herbaceous hosts in terrestrial habatats.

Sexual morph: Ascomata semi-immersed to erumpent, soli-

tary, scattered, broadly oblong, globose to subglobose, dark

brown to black, coriaceous, ostiolate. Ostiole single, central,

slot-like, with ostiolar canal filled with hyaline cells. Peridium

thick, 2-layered at the sides and at base, outer layer comprising

heavily pigmented, brown to black, thick-walled cells of

textura angularis; inner layer composed of hyaline, thin-

walled cells of textura prismatica; single layer of melanised,

occluded cells at apex. Hamathecium lacking

pseudoparaphyses.

Asci 8-spored, bitunicate, fissitunicate, cy-

lindrical to cylindric-clavate, short or bulbous pedicellate, api-

cally rounded, with a minute ocular chamber. Ascospores 1–2-

seriate, initially hyaline, becoming brown at maturity,

muriform, mostly ellipsoidal, 3–6transverselyseptate,with

1–4 longitudinal septa, narrowly rounded at both ends, with-

out a mucilaginous sheath, smooth-walled. Asexual morph:

Undetermined.

Fungal Diversity

Notes: A new genus Pseudostrickeria is introduced here to

accommodate Pseudostrickeria muriformis and

Pseudostrickeria ononidis based on morphology and phylo-

genetic analysis (Fig. 1). Muriform ascospores, the most im-

portant diagnostic character, are reported for the first time in

this family, in addition to the unusual lack of

pseudoparaphyses. Barr (1990a) included Strickeria in

Melanommataceae, while the unitunicate asci would suggest

this genus should be in Sordariomycetes

(Maharachchikumbura et al. 2015). In this study,

Fig. 25 Ohleria modesta (isotype). a, b Specimens and herbarium label.

c, d Appearance of gregarious ascomata on host surface. e Vertical hand-

cut section of ascoma. f Vertical hand-cut sections through peridium. g

Immature ascus. h–j Asci with ascospores. k Cellular pseudoparaphyses.

l–m Ascospores. o–p Ascospores breaking into two conical partspores.

Scale bars: c=500μm, d=200μm, e=50μm, f=20μm, g–k, o–p=10μm,

l–n=5μm

Fungal Diversity

P. muriformis (MFLUCC 13–0764) and P. ononidis

(MFLUCC 14–0949) cluster together to form a single clade

in Melanommataceae with 74 % ML BS (Fig. 1).

Pseudostrickeria muriformis is morphologically similar to

P. ononidis, while the former has ascomata with a carbona-

ceous peridium and 4–6transverselyseptate,2–4verticalsep-

tate ascospores; the latter has semi-immersed ascomata with a

reduced clypeus and 2–5 transversely septate, ascospores with

1–2 vertical septa. Thus, based on both morphological and

molecular characters these new collections are introduced as

new species, P. muriformis and P. ononidi.

Pseudostrickeria muriformis Wanasinghe, Q. Tian,

Camporesi & K.D. Hyde, sp. nov.

Index Fungorum number: IF551599 , FacesofFungi

number: FoF 01033, Fig. 26

Etymology: The specific epithet muriformis is based on the

ascospore septation.

Saprobic on dead upright stems of Origanum vulgare in

terrestrial habtats. Sexual morph: Ascomata 150–250μm

high, 250–400μm diam. (x̅=194.5×312.2μm, n=10), semi-

immersed and raising host tissue, to erumpent, sol itary,

scattered, broadly oblong, globose to subglobose, dark brown

Fig. 26 Pseudostrickeria

muriformis (holotype). a

Appearance of ascomata on host

surface. b Vertical hand-cut

section of ascoma. c Vertica l

hand-cut section through

peridium. d Immature ascus. e–g

Asci with ascospores. h–k

Ascospores. Scale bars:

b=100μm, c–g=20μm,

h–k=10μm

Fungal Diversity

to black, carbonaceous, ostiolate. Ostiole 25–50μm high, 40–

60μm diam. (x̅=38.2×50.1μm, n=5), single, centra l, slot-

like, black, smooth, with ostiolar canal filled with hyaline

cells. Peridium 5–15μm, thick at the base and 10–20μm wide

at sides, becoming carbonaceous and thicker above; 2-layered,

outer layer comprising heavily pigmented, thick-walled,

blackish to dark brown cells of textura angularis, inner layer

composed of hyaline, thin-walled cells of textura prismatica.

Hamathecium lacking pseudoparaphyses. Asci 90–110× 13–

17μm (x̅=101.8×15.1μm, n =40), 8-spored, bitunicate,

fissitunicate, cylindrical to cylindric-clavate, with short or bul-

bous pedicel, apically rounded, with a minute ocular chamber.

Ascospores 15–25×8–10μm (x̅=19.5×8.7μm, n=50), partly

overlapping 1–2-seriate at the apex, 1-seriate at the base, ini-

tially hyaline, becoming brown at maturity, muriform, mostly

ellipsoidal, with 4–6 transverse septa, and 2–4 longitudinal

septa, conical and narrowly rounded at the ends, without a

mucilaginous sheath. Asexual morph: Undetermined.

Culture characteristics: Colonies on PDA reaching 20–

25 mm diam at 25 °C after 21 d, spreading with sparse aerial

mycelium and lobed, feathery margins; surface olivaceous

grey in the centre, pale olivaceous-grey in the outer region;

in reverse olivaceous-grey in the centre, pale olivaceous-grey

in the outer region.

Material examined: ITALY, Provi nce of Forlì-Cesena,

Verghereto, Monte Fumaiolo, on dead upright stems of

Origanum v ulgare L. (

Lamiaceae), 22 August 2013, E.

Camporesi IT1420 (MFLU 15–1407, holotype; HKAS–

88735 isotype), ex-type living culture MFLUCC 13–0764=

KUMCC15-0062.

Pseudostrickeria ononidis Q. Tian, Wanasinghe,

Camporesi & K.D. Hyde, sp. nov.

Index Fungorum number: IF551600 , FacesofFungi

number: FoF01034, Figs. 27 and 28

Etymology: In refe rence to its occurrence on Ononis

spinosa, ononidis meaning Bof Ononis^.

Holotype:MFLU15–1410

Saprobic on dead stem of Ononis spinosa in terrestrial

habatats. Sexual morph: Ascomata 200–288μm high, 255–

300μm diam. (x̅=233.5×265.5μm, n=10), semi-immersed,

solitary, scattered, broadly oblong, dark brown to black, cori-

aceous, ostiolate. Ostiole 25–53μm high, 46–68μm diam. (x̅=

34.2×59.5μm, n=5), single, central, slot-like, black, smooth,

with ostiolar canal filled with hyaline cells. Peridium 8–18μm

wide at the base, 16–23

μm wide in sides, comprising 2 layers,

outer layer heavily pigmented, thick-walled, comprising

blackish to dark brown cells of textura angularis, inner layer

composed of hyaline, thin-walled, irregular cells of textura

prismatica. Hamathecium lacking pseudoparaphyses. Asci

87–120×14–17μm (x̅ =113×16μm, n =40), 8-spored,

bitunicate, fissitunicate, cylindrical to cylindric-clavate, short

or bulbous pedicellate, apically rounded, with a minute ocular

chamber. Ascospores 17–25×7–10μm (x̅=18.5×8.5μm, n=

50), partly overlapping 1–2-seriate at the apex, 1-seriate at

the base, muriform, mostly ellipsoidal, truncate at one side,

2–5 transversely septate, with 1–2 longitudinal septa, initially

hyaline, becoming brown at maturity, conical and narrowly

rounded at the ends, without a mucilagin ous sheath.

Asexual morph: Mycelium branched, septate, hyaline to pale

brown. Conidiophores 25–30(−36)×5–6μm (x̅=27×5.3μm,

n=10), hyaline, septate, cylindrical, formed from mycelium.

Conidiogenous cells holoblastic, basipetal, hyaline, smooth.

Conidia 7–12×6–7μm (x̅=8.8×6.3μm, n=10), aseptate, pale

brown to brown, eguttulate, globose to subglobose, truncate at

base, smooth-walled.

Culture characteristics: Colonies on PDA reaching 20–

25 mm diam at 25 °C after 21 d, surface umber with patches

of dirty white and iron-grey; with sparse to moderate aerial

mycelium and feathery margins; reverse iron.

Material examined: ITALY, Forlì-Cesena Province,

Verghereto, Sorgenti de l Savio, on dead stem o f Ononis

spinosa L. (Leguminosae), 10 March 2014, E. Camporesi

IT1760 (MFLU 15–1410, holotype;HKAS–88738, isotype),

ex-type living culture MFLUCC 14–0949=KUMCC15-

0061.

Sarimanas Matsumura, K. Hiray. & Kaz. Tanaka.

FacesofFungi number: FoF00496

Saprobic on twigs of various woody plants in terrestrial

environments. Sexual morph: Ascomata immersed, scattered

or gregarious, globose to subglobose, ostiolate. Neck short

papillate, without clypeus. Peridium composed of polygonal

thin-walled cells. Hamathecium elements cellular, numerous,

septate. Asci bitunicate, fissitunicate, cylindrical to ovoid, with

a short pedicel. Ascospores broadly fusiform with rounded

ends, 1-septate, hyaline, smooth-walled, with an entire gelat-

inous sheath. Asexual morph: Undetermined.

Type species: Sarimanas shirakamiense Matsumura, K.

Hiray. & Kaz. Tanaka, sp. nov., FacesofFungi number:

FoF00497

Notes: Sarimanas was established by Matsumura, K.

Hiray. & Kaz. Tanaka (2015) to accommodate

S. shirakamiense and S. pseudofluviatile. Morphologically,

Sarimanas is similar to Massarina, however, phylogenetic

analyses indicate that this genus has a close affinity to the

genera in Melanommataceae (Liu et al. 2015). In this study,

phylogenetic analysis of S. shirakamiense (MAFF 242969

and MAFF 244768) and S. pseudofluviatile (MAFF 239465)

based on four genes (LSU, SSU, RPB2 and EF-1α) was un-

dertaken (Fig. 1). This indicates that S. shirakamiense is sim-

ilar to S. pseudofluviatile and forms a well-supported clade

(83 % ML BS and 1.00 BY PP).

Xenostigmina Crous, Mycol. Mem. 21: 154 (1998)

FacesofFungi number: FoF 00781

Sarprobic on the surface of leaves in terrestrial habitats.

Conidiomata sporodochial, hypophyllous, black.

Conidiophores verruculose, brown, 1– 3-septate,

Fungal Diversity

subcylindrical, variously curved, branched, arising from the

upper cells of a suprastomata. Conidiogenous cells terminal

and intercalary, hyaline to light brown, verruculose,

subcylindrical, proliferating sympo dially. Conidia solitary,

broadly ellipsoidal to obclavate or subcylindrical, straight to

curved, obtuse at apex, truncate at base, muriform, with four to

eight horiz ontal a nd one t o three longitudinal septa,

verruculose.

Type species: Xenostigmina zilleri (A. F unk) Crous,

Mycol. Mem. 21: 155 (1998), Fig. 29

Notes: Xenostigmina was introduced by Crous (1998a, b)

to accommodate the type species Xenostigmina zilleri (A.

Fig. 27 Pseudostrickeria ononidis (holotype). a, b, c Appearance of

gregarious ascomata on the host surface. d Vertical section through

ascomata. e Vertical section through ostiole. f Vertical section through

peridium. g–j Asci with ascospores. Note the bitunicate asci with ocular

chamber. k–n Ascospores. Scale bars: d=100μm, e=20μm, c=100μm,

f–j=10μm, k–n=5μm

Fungal Diversity

Funk) Crous and for which Stigmina zilleri A. Funk is a syn-

onym. Currently, there are only two species in this genus (Kirk

et al. 2015), viz. X. wolfii Crous & Corlett and X. zilleri (A.

Funk) Crous. However, X. wolfii (Crous and Corlett 1998),

which is the asexual morph of Mycosphaerella stigmina-

platani, and a Pseudocercospora synanamorph, in thus ac-

commodated in Pseudocercospora (C rous et al . 2006).

Crous (1998a, b) revealed that Xenostigmina with its

Mycopappus synanamorph is distinct from Stigmina s. str.,

which has not been shown to reside in Pseudocercospora,

but appears to be related to Seifertia (Seifert et al. 2007)in

the Dothideomycetes (Crous et al. 2009). Subsequently,

Xenostigmina and Mycopappus grouped in

Phaeosphaeriaceae in the phylogenetic analysis by Crous

et al. (2013), whereas Phookamsak et al. (2014) excluded

the genus from Phaeosphaeriaceae and transferred it t o

Melanommataceae. The present multigene analysis indicated

that Xenostigmina zilleri (CBS 124108, CBS 115685, CBS

115686) is aligned with Mycopappus aceris (CBS 124109)

and formed a single clade in Melanommataceae with 93 %

MLBSand0.99BYPP.Xenostigmina is confirmed as

synanamorph of Mycopappus.Thus,Xenostigmina is retained

in Melanommataceae.

Excluded and doubtful genera

Based on phylogenetic analysis, we place Acrocordiopsis in

Aigialaceae, Beverwykella in Pleomassariaceae, while the

placement of Caryosporella is still uncertain in

Dothideomycetidae genus incertae sedis. Astrosphaeriella is

placed in Astrosphaeriellaceae

. We introduce a new genus

Thysanolaenae to accommodate two new taxon previously

published in Pseudotrichia. Sporidesmiella is also transferred

to Roussoellaceae.

Fig. 28 Colony cultures of

Pseudostrickeria ononis a

Colonies on MEA. b, c, e, f

Conidiophores with conidia

from culture. d Vegetative

hyphae in culture. Scale bars:

b, c, e, f=10 mm, d=5μm

Fig. 29 Xenostigmina zilleri from DAVFP 23272 (re-drawn from Crous

1998). Conidiophores and conidia. Scale bars=10μm

Fungal Diversity

Aigialaceae Suetrong, Sakay.et al., in Suetrong et al., Stud.

Mycol. 64: 166 (2010)

Notes: Aigial ace ae was introduced by Suetrong et al.

(2009) to include three genera, Aigialus, Ascocratera and

Rimora which were previously assigned to Massarinaceae

(Lumbsch and Huhndorf 2010). Zhang et al. (2012) noted that

despite the wide range of morphological variation between

members of Aigialaceae, the family is strongly supported

(75/1.00 ML/BS) in phylogenetic analyses of four loci

(LSU, SSU, RPB2 and EF-1α)inPleosporales as shown in

Fig. 1. In this study, Acrocordiopsis are included as sister

groups of Aigialus in Aigialaceae.

Acrocordiopsis Borse & K.D. Hyde, Mycotaxon 34(2):

535 (1989)

FacesofFungi number: FoF 00786

Saprobic on decaying mangroves in marine habitats.

Sexual morph: Ascomata solitary or gregarious, immersed,

beneath a raised dark brown to black pseudoclypeus, conical

to subglobose, black, ostiolate. Peridium 2-layered.

Hamathecium comprising filiform, persistent, branched, long,

colourless, trabeculate pseudoparaphyses, encircling the asci.

Asci 8-spored, bitunicate, fissitunicate, cylindrical to cylindro-

clavate, with a short pedicel, apically rounded and thickened,

with an ocular chamber. Ascospores uni-seriate, hyaline, turn-

ing brown at maturity, ovoidal or ellipsoidal, rounded at both

ends, or tapering towards subacute ends, 1-septate, constricted

at the septum, with two oil globules in each cell, smooth-

walled. Asexual morph: Undetermined.

Type species: Acrocordiopsis patilii Borse & K.D. Hyde

Notes: Borse and Hyde (1989) introduced the monotypic

genus Acrocordiopsis to accommodate Acrocordiopsis patilii.

The characteristics of which, indicated that Acrocordiopsis

should be assigned to Melanommatales (sensu Barr 1983).

Acrocordiopsis has affinities with Acrocordia, a lichen genus

with asci possessing a meniscus that stains in Congo Red

(Alias et al. 1999), while Acrocor diopsis has unstaining, api-

cally thickened asci. Acrocordiopsis also shares similarities

with Astrosphaeriella

,whileAstrosphaeriella has elongate-

fusoid ascospores with a gelatinous sheath. Jones et al.

(2009)assignedAcrocordiopsis to Melanommataceae based

on morphological characters. Phylogenetic analysis showed

that Acrocordiopsis patilii nested within Salsugineaceae

(Suetrong et al. 2009; Hyde et al. 2013). In this study, the

phylogenetic analyses based on a combined dataset of LSU,

SSU, RPB2 and EF-1α sequences includes Acrocordiopsis

patilii (BCC 28166 and BCC 28167), which clusters with

Salsuginea ramicola (KT 2597.1) in Aigialaceae. Thus

Acrocordiopsis canbeexcludedfromMelanommataceae.

Acrocordiopsis patilii Borse & K.D. Hyde, Mycotaxon

34(2): 536 (1989)

FacesofFungi number: FoF 00787, Fig. 30

Saprobic on decaying wood submerged in brackish waters

in mangroves. Sexual morph: Ascomata 1200–1800μm

high×(1500–)1800–3200μm diam. (x̅=1550×2850μm, n=

10), scattered or gregarious, immersed, erumpent to nearly

superficial, beneath a raised dark brown to black

pseudoclypeus, conical or subglobose, with a flattened base,

black, carbonaceous, extremely difficult to cut, ostiolate.

Ostiole single, papillate, with a centre ostiole, cone-shaped,

brown to black, shiny, smooth. Peridium up to 650μm thick

near the apex, 250–310μm thinner at the base, two-layered,

outer cells pseudoparenchymatous with small melanised,

thick-walled cells of textura epidermoidea, walls 6–11μm

thick; inner cells lighter, comprised of hyaline to pale brown,

irregular, elongate, thin-walled cells of textura angularis.

Hamat hecium of 1–1.5μm broad, dense, long, trabeculate

pseudoparaphyses, embedded in mucilage, hyaline, anasto-

mosing and sparsely septate. Asci 130–240×13–15(−18) μm

(x̅=220×14μm, n=10), 8-spored, bitunicate, fissitunicate, cy-

lindrical to cylindro-clavate, with a short pedicel, up to

28(−40) μm long, apically rounded, with a large and conspic-

uous ocular chamber. Ascospores (14–)17–25×12–17(−20)

μm (x̅=21×14μm, n=10), uni-seriate or partially overlapping

uni-seriate, hyaline, becoming pale brown when mature,

ovoid or ellipsoidal, rounded at both ends, or tapering toward

subacute ends, 1-septate, constricted at the septum, with two

oil guttules in each cell, smooth-walled. Asexual morph:

Undetermined.

Material examined: INDIA, Ind ian Ocean, Malvan,

Maharashtra, on intertidal wood of Avicennia alba Blume

(Acanthac eae), 30 October 1981, E.B.G. Jones (IMI

297769, holotype).

Astrosphaeriellaceae Phookamsak & K.D. Hyde, Fungal

Divers. In press

Notes: Astrosphaeriellaceae is introduced to accommodate

Astrosphaeriella and Pteridiospora by Phookamsak et al.

(2015) based on phylogenetic analyses. It is characterized by

a unique character of carbonaceous ascomata and trabeculate

pseudoparaphyses.

Astrosphaeriella Syd. & P. Syd., Annls mycol. 11(3): 260

(1913)

Notes: The genus and related genera have been dealt with

in a concurrent paper by Phookamsak et al. (2015). The genus

does not belong in Melanommataceae and the new families

Astrosphaeriellaceae and Pseudoastrosphaeriellaceae were

introduced to accommodate its species.

Pleomassariaceae M.E. Barr, M ycologia 71(5): 949

(1979)

Notes:

Pleomassariaceae was introduced by Barr (1979)as

a new family including Pleomassaria, Splanchnonema and

Asteromassaria. Currently the family comprises

Lichenopyrenis, Splanchnonema, Peridiothelia and

Pleomassaria (Zhang et al. 2012) which has been synony-

mized under Melanommataceae (Zhang et al. 2009a). The

asexual states of the Pleomassariaceae are mostly

coelomycetous. Zhang et al. (2012) reinstated

Fungal Diversity

Pleomassariaceae as a separate family in the order

Pleosporales based on phylogenetic analysis and included

the genera Lichenopyrenis, Splanchnonema, Peridiothelia

and Pleomassaria.

Beverwykella Tubaki, Trans.Mycol. Soc. Japan 16(2): 138

(1975)

FacesofFungi number: FoF 00762

Sexual morph: Undetermined. Asexual morph:

Conidiophores macronematous, mononematous, light fuscous

at maturity, septate, unbranched, ere ct, s traight to sligh tly

flexuous. Conidiogenous cells integrated, holoblastic, termi-

nal. Conidia hyaline, fuscous when mature, lenticular, hori-

zontally flattened; viewed from above, oval to almost rectan-

gular (with rounded edges); consisting of an internal, strictly

dichotomous branching system and an outer distinctly clath-

rate layer, which is produced by eight outermost branches

(four on upper side, four on lower side of a conidium) of the

internal branching system (van Beverwijk 1954; Voglmayr

and Delgado-Rodríguez 2003).

Type species: Beverwykella pulmonaria (Beverw.) Tubaki

Fig. 30 Acrocordiopsis patilii

(holotype). a, b Specimens and

herbarium label. c–f Appearance

of ascomata on host surface. g–h

Hand-cut section of ascoma. i

Hamathecium. j–m Asci with

ascospores. n–q Ascospores.

Scale bars: c=1000μm,

d–f=500μm, g–h=100μm,

j–m=10μm, i, n–q=5μm

Fungal Diversity

Notes: Beverwykella was introduced by Tubaki (1975). The

type sp ecies Beverwykella pulmonaria was originally de-

scribed as Papulaspora pulmonaria (Beverwijk 1954), which

is a very common aeroaquatic species throughout the temper-

ate to subtro pical Northern Hemisphere (Voglmayr and

Delgado-Rodríguez 2003). Beverwykella is characterized by

lenticular, black, multicellular conidia and a vertical main

branch and a series of mainly opposite side branches

(Beverwijk 1954, Tubaki 1975; Michaelides and Kendrick

1982). The type species lacks molecular data, there has been

few studies of B. pulmonaria, and the holotype was not ex-

amined. Currently, there are three species listed in

Beverwykella (Kirk et al. 2015), viz. B. cerebriformis

Nawawi & Kuthub.; B. clathrata Vo gl m ay r a nd

B. pulmonaria (Beverw.) Tubaki. Based on LSU and RPB2

seuqence data of CBS 283.53, Zhang et al. (2009b) studied the

phylogenetic relationship of B. pulmonaria, which proved to

be the asexual morph of Melanomma pulvis-pyrius (Chesters

1938; Sivanesan 1984). Gruyter et al. (2013) indicated that

B. pulmonaria is close to Aposphaeria corallinolutea, but

should not be the asexual morph of Melanomma pulvis-pyrius.

In the present study, we collected fresh B. pulmonaria from

England, and assigned as reference specimens, which clus-

tered with B. pulmonaria (CBS 283.53) with high support

(100 % ML BS and 1.00 BY PP) but represent a single clade

in Pleomassariaceae and thus Beverwykella is excluded from

Melanommataceae.

Beverwykella pulmonaria (Beverw.) Tubaki, Trans.

Mycol. Soc. Japan 16(2): 139 (1975).

Synonymy:

≡ Papulaspora pulmonaria Beverw., Antonie van

Leeuwenhoek 20: 11 (1954)

FacesofFungi number: FoF 00762, Figs. 31 and 32

Saprobic on Fagus sylvatica L. seeds. Asexual morph:

Colonies effuse, brown, and inconspicuous. Mycelium partly

superficial, partly immersed in the substratum, composed of

smooth, branched, septate, brown hyphae 1.5–3μm

wide.

Conidiophores macronematous, mononematous, light fuscous

at maturity, septate, unbranched, erect, straight to slightly flex-

uous, 35–50μm long and 2.5–3.5μm wide, not widened at the

base. Conidiogenous cells integrated, holoblastic, terminal.

Conidia 25–28μm high, 40–60μm diam, young hyaline, then

becoming brown, Mature conidium 77–92μm high, wide 70–

87μm, fuscous, lenticular, horizontally flattened, from above

oval to almost rectangular (with rounded edges); consisting of

an internal, strictly dichotomous branching system and an

outer distinctly clathrate layer, which is produced by eight

outermost branches (four on upper side, four on lower side

of a conidium) of the internal branching system (Beverwijk

1954; Voglmayr and Delgado-Rodríguez 2003). Sexual

morph: Undetermined.

Culture characteristics: Spores germinated on Malt Extract

Agar medium. Colonies on MEA appressed, circular, cream at

first, then becoming dark brown and raised mycelium in the

centre, reverse brown, with slow growth.

Material examined: UK, Bishops Waltham, Hampshire,

on Fagus sylvatica (Fagaceae), 29 September 2014, G. Jones

GJ 062 (MFLU 15–1398, HKAS, reference specimen de-

sign ed here), ex-type living culture MFUCC 15–0183=

KUMCC.

Caryosporella Kohlm., Proc. Indian Acad. Sci., Pl. Sci.

94(2–3): 355 (1985)

FacesofFungi number: FoF 00768

Saprobic on decaying mangrove wood in marine habitats.

Sexual morph: Ascomata superficial, seated on a thin, black

stroma, scattered or gregarious, sometimes densely gregari-

ous, subglobose, carbonac eous, periph ysate, ost iolate.

Ostiole with a distinct papilla, black. Peridium thick, 3-lay-

ered, outside cells yellow, forming a textura epidermoidea,

brown in the middle, also forming a textura epidermoidea,

hyaline cells inside and forming a textura prismatica.

Hamathecium of dense, trabeculate pseudoparaphyses,

branched and anastomosing above asci, embedded in a gelat-

inous matrix. Asci 8-spored, bitunicate, fissitunicate, cylindri-

cal, with a long pedicel, thick-walled, apically rounded with

an ocular chamber. Ascospores ellipsoidal to broadly fusoid,

both ends narrowing abruptly to a hyaline germ pore, 1-sep-

tate, thick-walled, deep reddish brown to dark brown, indis-

tinctly verruculose. Asexual mo rph: suspected spe rmatia

(Kohlmeyer 1985).

Type species: Caryosporella rhizophorae Kohlm.

Notes: Caryosporella was i ntroduced by Kohlmeyer

(1985) based on the obligate marine fungus, C. rhizophorae,

from dead wood of intertidal roots and branches of

Rhizophora mangle from Belize (Central America). The ge-

nus is characterized by superficial ascomata, a 3-layered pe-

ridium, trabeculate pseudoparaphyses and brown, 1-septate

ascospores (Kohlmeyer 1985). Caryosporella was compared

with Caryospora, previously assigned to Massariaceae,but

subsequently, Caryosporella was transfered to

Melanommataceae (Eriksson 2006; Lumbsch and Huhndorf

2007, 2010;Hydeetal.2013; Wijiyawardene et al. 2014).

Suetrong et al. (2009) demonstrated that a single isolate of

C. rhizophorae is related to Lineolata rhizophorae (Kohlm.

& E. Kohlm.) Kohlm. & Volkm.-Kohlm and did not cluster in

the Pleosporales based on LSU, SSU, RPB2 and EF-1α anal-

ysis, so was placed in the Dothideomycetidae incertae sedis.

Caryosporella is monotypic and has been placed in

Melanomm atacea e by Zhang et al. (2012), Hyde et al.

(2013), and Wijiyawardene et al. (2014). However, the famil-

ial placement of Caryosporella rhizophorae is acknowledged

to be uncertain and it may not belong to the Pleosporales

(Zhang et al. 2012). In this study, based on four genes (LSU,

SSU, RPB2 and EF-1α), C. rhizophorae (JK 5302A) forms a

single clade but does not cluster i n Melanommataceae.

Therefore it i s proposed that Caryosporella should be

Fungal Diversity

excluded from Melanommataceae and placed in

Dothideomycetes, genus incertae sedis.

Caryosporella rhizophorae Kohlm., Proc. Indian Acad.

Sci., Pl. Sci. 94(2–3): 356 (1985)

FacesofFungi number: FoF 00769, Fig. 33

Saprobic on dead wood of intermittently submerged roots

or branches of Rhizophora mangle L.. Sexual morph:

Ascomata 800–1000μm high×900–1100μm diam (x̅=927×

990μm, n=10), superficial with a flattened base, seated on a

thin black stroma, scattered or densely gregarious,

subglobose, carbonaceous, periphysate, ostiolate. Ostiole

with a di stinct papilla, black. Peridium thicker at the

apex, up to 200μm wide , thinne r at both sides, 120–

150μm thick, 3-layered, outer cells yellow, thick-walled

cells of textura epidermoidea, brown in the middle layer

with irregular cells of textura epidermoidea,innerlayer

composed of hyaline, thin-walled cells of textura

prismatica. Hamathecium of dense, (1–)1.5–2μm wide,

trabeculate pseudoparaphyses, long, branched and anas-

tomosing above asci, ernbedded in a gelatinous matrix.

Asci 220–27 0(−300)×14–17(−20) μm (x̅=246×16.5μ m,

n =10), 8-spored, bitunicate, fissitunicate, cylindrical,

with a long pedicel, thick-walled, apically rounded, with

an ocular chamber.

Ascospores 24–28(−30)×8–13 μm

(x̅=27×11.5μm, n=10), uni-seriate or partially overlap-

ping uni-seriate, ellipsoidal to broadly fusoid, sometimes

curved, both ends narrowing abruptly to a hyaline germ

pore, 1-septate, slightly constricted at the septum, thick-

walled, deep reddish brown to dark brown, indistinctly

verruculose. Asexual morph: Undeterm ined.

Fig. 31 Papulaspora

pulmonaria (holotype,redrawn

from Beverwijk 1954) a Erect

young conidiophore. b, c, e

Conidial initial showing branch

system and sphaerical cells

terminating the dichotomous

divisions. d, g Premature

conidium becoming the compact

outermost cell layer at the end of

dichotomous branching. f

Sphaerical cells of conidium. h

Section of mature conidium. I

Mature conidia. Scale bars:

a–i=10μm

Fungal Diversity

Material examined: BELIZE, Twin Cays, on dead wood

of intermittently submerged roots or branches of Rhizophora

mangle (Rhizophoraceae), 3 April 1983, J. Kohlmeyer

(4532a, holotype).

Didymosphaeriaceae Munk, Dansk bot. Ark. 15(no. 2):

128 (1953)

Basionym:

= Montagnulaceae M.E. Barr, Mycotaxon 77: 194 (2001)

Notes:ThefamilyDidymosphaeriaceae was introduced by

Munk A. (1953) which is characterized by 1-septate asco-

spores and trabeculate pseudoparaphyses (Aptroot 1995;

Ariyawansa et al. 2014c). Ariyawansa et al. (2014d)included

Didymosphaeria and genera from Montagnulaceae in

Didymosphaeriaceae. They synonymise Montagnulaceae

under Didymosphaeriaceae as Didymosphaeriaceae is oldest

name.

Pseudotrichia Kirschst., Annls mycol. 37(1/2): 125 (1939)

FacesofFungi number: FoF 00784

Saprobic in terrestrial or rarely aquatic habitats. Sexual

morph: Ascomata, solitary or scattered, initially immersed,

erumpent to nearly superficial, globose to pyriform, carbona-

ceous or coriaceous, ostiolate. Ostiole single, central, papil-

late. Peridium two-layered, comprising of thick-walled cells

of text ura angularis, outer layer heavily pigmented, small

cells, dark brown to black. Hamathecium numerous, dense,

hyaline, septate, narrow, unbranched, cellular

pseudoparaphyses, embedded in mucilage. Asci 8-spored,

bitunicate, fissitunicate, elongate clavate to sub-cylindrical,

Fig. 32 Beverwykella pulmonaria (reference specimen). a Host Fagus sylvatica cupule b, c Colonies on the substrate d–h Single mature bulbil. i Cells

of mature bulbil. Scale bars: b=100μm, c=200μm, d–h=30μm, i=10μm

Fungal Diversity

pedicellate, apically rounded with a minute ocular chamber.

Asco spores uniseriate or bitunicate, hyaline to rarely pale

brown, fusoid, 1-septate or with multiple septa, constricted

at the septum, with a large guttule in each cell, smooth-walled,

thick-walled. Asexual morph: Undetermined.

Type species: Pseudotrichia stromatophila Kirschst.

Notes: Pseudotrichia was introduced by Kirschstein (1939)

as a monotypic genus to accommodate P. stromatophila.

There are 13 species assigned to this genus (Kirk et al.

2015). The characters of ascospores, such as size and the

number of septa, are highly varied in this genus (Barr 1984;

Fallah and Shearer 200 1). Thambugala et al. (2014)re-

examined the type species P. stromatophila and accommodat-

ed Pseudotrichia in Montagnulaceae based on ascomatal and

peridial characters, cellular, septate, unbranched

pseudoparaphyses and asci with long pedicels. Liu et al.

(2015) introduced two new species P. rubriostiolata and

P. thailandica which clustered in Melanommataceae based

on multigene analysis. In the present study, P. rubriostiolata

and P. thailandica formed a single clade between

Fig. 33 Caryosporella

rhizophorae (holotype). a

Superficial ascomata on the

surface of host. b Vertical section

through ascoma. c Hand-cut

section through peridium. d, e, g–

j Asci with ascospores, note the

fissitunicate state of some asci. f

Trabeculate pseudoparaphyses.

k–n Ascospores. Scale bars:

a=1000μm, b=100μm,

c=50μm, d–e, g–j=10μm,

f, k–n=5μm

Fungal Diversity

Roussoellaceae and Aigialaceae, and is morphologically dis-

tinct from P. stromatophila. Pseudotrichia rubriostiolata and

P. thailandica have broad, clavate asci with short, foot-like

pedicels, while P. stromatophila has elongate, clavate to sub-

cylindrical asci with a long pedicel. The peridia of the species

also differ. Thus, we introduce a new genus Thysanolaenae to

accommodate these two species. Molecular data for the type

species, P. stromatophila is not available in GenBank, thus we

follow Thambugala et al. (2014), and exclude Pseudotrichia

from Melanommataceae and retain it in Didymosphaeriaceae

(Basionym: Montagnulaceae).

Thysanolaenae Q. Tian & K.D. Hyde, gen. nov.

Index Fungorum number: IF551601 , FacesofFungi

number: FoF 01271

Etymology: In reference to the host Thysanolaena maxima

(Poaceae).

Saprobic on Thysanolaena maxima. Sexual morph:

Ascomata solitary, scattered, immersed to semi-immersed

erumpent with neck, with reddish pigment surrounding the

ostiole, globose to subglobose, ostiole central, with short pa-

pilla. Peridium thin-walled, comprising 2–3 layers.

Hamathecium composed of dense, narrow cellular

pseudoparaphyses, anastomosing above the asci, embedded

in a hyaline gelatinous matrix. Asci 8-spored, bitunicate, cla-

vate, pedicellate, with a short foot-like pedicel, apically round-

ed, with an ocular chamber. Ascospores 1–3-seriate, fusiform,

slightly curved, hyaline, becoming light brown when gemi-

nated, 1-septate, constricted a t the septa, smooth-walled,

surrounded by distinct mucilaginous sheath. Asexual morph:

Undetermined.

Type species: Thysanolaenae rubriostiolata (Phookamsak

& K.D. Hyde) Q. Tian & K.D. Hyde.

Notes: A new genus Thysanolaenae is introduced here to

accommodate T. rubriostiolata and T. thailandica which have

been placed in Pseudotrichia based on phylogenetic analysis

(Fig. 1). These two species share similarities, while the

ascomata, asci and ascospores are smaller in

P. rubriostiolata than in P. thailandica.

Thysanolaenae rubriostiolata (Phookamsak & K.D.

Hyde) Q. Tian, & K.D. Hyde.

Basionym:

= Pseudotrichia rubriostiolata Phookamsak & K.D. Hyde,

in Liu et al., Fungal Diversity: 10.1007/s13225-015-0324-y,

[123] (2015)

Index Fungorum number:IF551577

Description see Liu et al. (2015)

Thysanolaenae thailandica (Phookamsak & K.D. Hyde)

Q. Tian, & K.D. Hyde.

Basionym:

= Pseudotrichia thailandica Phookamsak & K.D. Hyde, in

Liu et al., Fungal D iversity: 10.1007/s132 25-015-0 324- y,

[123] (2015)

Index Fungorum number:IF551602

Description see Liu et al. (2015)(Table4)

Roussoellaceae J.K. Liu, R. Phoo kamsak, D.Q. Dai &

K.D. Hyde, Fungal Diversity 181: 7 (2014)

Notes: Roussoellaceae was introduced by Liu et al. (2014)

to accommodate Roussoella, Roussoellopsis and

Neoroussoella. This family is characterized as having semi-

immersed to immersed, clypeate ascostromata containing tra-

beculate pseudopara physes long cylindric al to clavate

bitunicate asci and 2-celled ornamented ascospores. All the

strains of Roussoellaceae formed a well-supported clade with-

in the order Pleosporales.

Sporidesmiella P.M. Kirk, Trans. Br. mycol. Soc. 79(3):

479 (1982)

FacesofFungi number: FoF 00790

Saprobic on woody host. Sexual morph: Undetermined.

Asexual mor ph: Mycelium partly superficial, partly im-

mersed in the substratum, comprising branched, septate, pale

brown to brown hyphae. Conidiophores macronematous,

mononematous, solitary, straight or slightly flexuous, septate,

brown to dark b rown. Conidiogenous cells holoblastic,

monoblastic, integrated, terminal, proliferating percerrently,

rarely sympodially. Conidia seceding schizolytically, cylindri-

cal, narrowly clavate, obovoid to broadly obovoid or cunei-

form, truncate at the base, round at the apex, 1–5-distoseptate,

with reduced cell lumina, pale olivaceous to olivaceous brown

or brown.

Type species: Sporidesmiella claviformis P.M. Kirk

Table 4 Difference between Pseudotrichia speices discussed in this study

Species Size Septation Ascospore colour Host References

Ascospore (μm)Asci(μm)Ascomata(μm)

Pseudotrichia

guatopoensis

44–52×6.5–8175–200×

22–27

450–550×500–

550

3–5 hyaline Woody twig Huhndorf 1994

Thysanolaenae

rubriostiolata

25–27×4–570–80×15–

140–220×150–

190

1–3 hyaline to light

brown

Thysanolaena maxima

(Poaceae)

This study

Pseudotrichia

stromatophila

35–40×7–12 135–200×

13–22

500–750×520–

710

1–3 hyaline to pale

brown

Diatrypella (Diatrypaceae) Kirschstein 1939

Thysanolaenae

thailandica

28–32×4–580–100×

16–18

140–180×165–

310

1–5 hyaline to

subhyaline

Thysanolaena maxima

(Poaceae)

This study

Fungal Diversity

Notes: Sporidesmiella was established by Kirk (1982)to

accommodate S. clavifo rmis Kirk (type species) and

S. longissima Kirk. Currently, Sporidesmiella contains 34 val-

id taxa (Kirk et al. 2015). In this study, we suggest

Sporidesmiella is polyphyletic with species clustering in two

clades. Sporidesmiella fusiformis (HKUCC 10831) aligns in

the family Montagnulaceae with 97 % ML BS and 1.00 BY

PP and Sporidesmiella minima (CBS 524.50) cluster in

Roussoellaceae. Thus, we exclude Sporidesmiella from

Melanommataceae.

Sporidesmiella claviformis P.M. Kirk, Trans. Br. mycol.

Soc. 79(3): 479 (1982)

FacesofFungi number: FoF 00791, Fig. 34

Saprobic on twigs of Ulex europaeus L.. Asexual morph:

Colonies hairy, sparse, brown to dark brown, on the surface of

Ulex europaeus. Mycelium superficial, partly immersed in the

Fig. 34 Sporidesmiella

claviformis (holotype). a, b

Specimens and herbarium label of

Sporidesmiella claviformis. c, d

Superficial pale brown

conidiophores. e–g

Conidiophores. h, i Conidia. j, k

Sporidesmiella claviformis (re-

drawn from Kirk 1982). Scale

bars: d–i=5μm. j–k=10μm

Fungal Diversity

substratum, comprising branched, septate, pale brown to

brown hyphae. Conidiophores 45–92(−158)×2–4.5μm (x̅=

74×2.8μm, n=20), solitary, macronematous, mononematous,

brown to dark brown, straight or slightly flexuous, septate,

swollen at the base, up to 12μm wide. Conidiogenous cells

integrated, holoblastic, monoblastic, terminal, proliferating

percurrently, rarely sympodially, cylindrical. Conidia 14–

21×5.8–8μm (x̅=18.5×6.5μm, n=10), solitary, seceding

schizolytically, pale olivaceous to olivaceous brown or brown,

cylindrical, narrowly clavate, obovoid to broadly obovoid or

cuneiform, truncate at the base, round at the apex, 2-

distoseptate, with reduced cell lumina, smooth-walled.

Sexual morph: Undetermined.

Material examined: UNITED KINGDOM, England,

Surrey, on twigs of Ulex europaeus (Fabac eae), 7 April

1980, P. M. Kirk 542 (IMI 246982, holotype).

Discussion

Melanommataceae is an interesting family with many species,

but phylogenetica lly poorly studied. Re collection of fresh

specimens and epitypification with molecular sequence data

are important to confirm the correct classification of taxa of

Melanommataceae. Trabeculate pseudoparaphyses were con-

sidered to be an important character at the ordinal level in

Pleosporales and Melannomatales (Barr 19 83), but Liew

et al. (2000) showed that this classification was not supported

at the molecular level. Trabeculate pseudoparaphyses are usu-

ally considered as narrower tha n 1μm,asinthecaseof

melanommataceous taxa (Hyde et al. 2000). In the present

study, taxa of Melanommataceae have narrow

pseudoparaphyses ranging from 1 to 2μm diam., usually anas-

tomosing and branching between and above the asci and em-

bedded in a gelatinous matrix. They are, however, not strictly

trabeculate pseudoparaphyses as in species of

Astrosphaeriella (see Phookamsak et al. 2015), which are

generally less than 1μ m diam. In some cases

pseudoparaphyses in Melannomataceae are lacking, such as

in the genera Pseudostrickeria and Muriformistrickeria.Care

must therefore be taking in the interpretation of trabeculate

versus cellular pseudoparaphyses, as they are difficult to dis-

tinguish and their distinction is not always supported by mo-

lecular data.

Acknowledgments MFLU grant number 56101020032 is thanked for

supporting studies on Dothideomycetes. We are also grateful to the Mush-

room Research Foundation, Chiang Rai, Thailand for supporting this

research. 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 In-

stitute of Botany. Jian-Chu Xu would like to thank the CGIAR Research

Program 6: Forest, Trees and Agroforestry, for partially funding this

work. Qing Tian and Putaruk Chomnunti thank the National Research

Council of Thailand (grant for Dothideomycetes No: 58201020010) for

supporting studies on Dothideomycetes. Dr. Shaun Pennycook is grate-

fully thanked for checking and revising the Latin name of new taxa. The

curators fro m G, K, S, NY, HKU and IFRD herbaria are gratefully

thanked for loaning specimens and type information. Rungtiwa

Phookamsak and Hongyan Su are thanked for providing the sequences

of Pseudotrichia species, Byssosphaeria musae and Phragmocephala

species.

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An overview of Melanommataceae (Pleosporales, Dothideomycetes): Current insight into the host associations and geographical distribution with some interesting novel additions from plant litter

Article

Full-text available

Jun 2024

Melanommataceous species exhibit high diversity with a cosmopolitan distribution worldwide and show a prominent saprobic lifestyle. In this study, we explored five saprobic species collected from plant litter substrates from terrestrial habitats in China and Thailand. A combination of morphological characteristics and multi-locus phylogenetic analyses was used to determine their taxonomic classifications. Maximum Likelihood and Bayesian Inference analyses of combined LSU, SSU, ITS and tef1-α sequence data were used to clarify the phylogenetic affinities of the species. Byssosphaeria poaceicola and Herpotrichia zingiberacearum are introduced as new species, while three new host records, Bertiella fici, By. siamensis and Melanomma populicola are also reported from litter of Cinnamomum verum, Citrus trifoliata and Fagus sylvatica, respectively. Yet, despite the rising interest in the melanommataceous species, there is a considerable gap in knowledge on their host associations and geographical distributions. Consequently, we compiled the host-species associations and geographical distributions of all the so far known melanommataceous species.

The rare fungicolous fungus Pseudotrichia mutabilis (Pers.) Wehm.: new data and a historical perspective

Article

Oct 2022

Olexander Akulov

The paper reports two new records of a rare fungicolous fungus Pseudotrichia mutabilis (Melanommataceae, Pleosporales) from Ukraine. Both specimens were collected within protected areas in association with two xylariaceous species, Hypoxylon crocopeplum and Rosellinia corticium. The paper provides descriptions, nomenclatural data, and original illustrations of the reported species. Information on substrate specialization and nutritional strategies of Pseudotrichia mutabilis, as well as general distribution of the species and known localities in Ukraine are specified and summarized.

Mycosphere notes 325-344 -Novel species and records of fungal taxa from around the world

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Full-text available

Jan 2021

Fungi have a worldwide distribution, however, less than 10% of the estimated species have

Two new species and a new host record of Pleosporales (Dothideomycetes) from palm ( Arecaceae ) in Guangdong Province, China

Article

Full-text available

Oct 2023

Pleomorphic Dematiomelanomma yunnanense gen. et sp. nov. (Ascomycota, Melanommataceae) from grassland vegetation in Yunnan, China

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Full-text available

Jul 2023

During a survey of microfungi associated with grasslands and related vegetation types from Yunnan Province in China, various ascomycetous and coelomycetous fungi were isolated. This study reports the discovery of four strains of ascomycetous and coelomycetous fungi from dead stalks of Hypericum monogynum L. (Hypericaceae) and Rubus parvifolius L. (Rosaceae) in the Zhaotong region of Yunnan Province, China. The isolates were characterized using multi-locus phylogenetic analyses and were found to represent a new monophyletic lineage in Melanommataceae (Pleosporales, Dothideomycetes). This new clade was named as Dematiomelanomma yunnanense gen. et sp. nov. which consists of both sexual and asexual morphs. The sexual morph is characterized by globose to subglobose ascomata with a central ostiole, cylindrical asci with a pedicel and ocular chamber, and muriform, ellipsoidal to fusiform ascospores. The asexual morph has synanamorphs including both brown, muriform macroconidia and hyaline, round to oblong or ellipsoidal microconidia. These findings contribute to the understanding of fungal diversity in grasslands and related vegetation types in Yunnan Province, China.

Development of PCR-Multiplex Assays for Identification of the Herpotrichiellaceae Family and Agents Causing Chromoblastomycosis

Preprint

Full-text available

Mar 2024

The Herpotrichiellaceae family is an important group of dematiaceous filamentous fungi, associated with a variety of pathogenic fungal species causing Chromoblastomycosis (CBM) and Phaeohyphomycosis (PHM), both with polymorphic clinical manifestations and worldwide incidence. Currently, the identification of this family and determination of the causative agent is challenging due to the subjectivity of morphological identification methods, necessitating the use of molecular techniques to complement diagnosis. In this context, genetic sequencing of the Internal Transcribed Spacer (ITS) has become the norm due to a lack of alternative molecular tools for identifying these agents. Therefore, this study aimed to develop PCR-Multiplex methodologies to address this gap. Sequences from the ITS and Large subunit (LSU) of ribosomal DNA were used, and after manual curation and in vitro analyses, primers were synthesized for the identification of the aforementioned targets. The primers were optimized and validated in vitro, resulting in two PCR-Multiplex methodologies: one for identifying the Herpotrichiellaceae family and the bantiana clade, and another for determining the species Fonsecaea pedrosoi and Fonsecaea monophora. Ultimately, the assays developed in this study aim to complement other identification approaches for these agents, reducing the need for sequencing, improving the management of these infections, and enhancing the accuracy of epidemiological information.

Palm Fungi and Their Key Role in Biodiversity Surveys: A Review

Article

Full-text available

Nov 2023
J. Fungi

Over the past three decades, a wealth of studies has shown that palm trees (Arecaceae) are a diverse habitat with intense fungal colonisation, making them an important substratum to explore fungal diversity. Palm trees are perennial, monocotyledonous plants mainly restricted to the tropics that include economically important crops and highly valued ornamental plants worldwide. The extensive research conducted in Southeast Asia and Australasia indicates that palm fungi are undoubtedly a taxonomically diverse assemblage from which a remarkable number of new species is continuously being reported. Despite this wealth of data, no recent comprehensive review on palm fungi exists to date. In this regard, we present here a historical account and discussion of the research on the palm fungi to reflect on their importance as a diverse and understudied assemblage. The taxonomic structure of palm fungi is also outlined, along with comments on the need for further studies to place them within modern DNA sequence-based classifications. Palm trees can be considered model plants for studying fungal biodiversity and, therefore, the key role of palm fungi in biodiversity surveys is discussed. The close association and intrinsic relationship between palm hosts and palm fungi, coupled with a high fungal diversity, suggest that the diversity of palm fungi is still far from being fully understood. The figures suggested in the literature for the diversity of palm fungi have been revisited and updated here. As a result, it is estimated that there are about 76,000 species of palm fungi worldwide, of which more than 2500 are currently known. This review emphasises that research on palm fungi may provide answers to a number of current fungal biodiversity challenges.

Ascomycetes from karst landscapes of Guizhou Province, China

Article

Full-text available

Oct 2023

This study documents the morphology and phylogeny of ascomycetes collected from karst landscapes of Guizhou Province, China. Based on morphological characteristics in conjunction with DNA sequence data, 70 species are identified and distributed in two classes (Dothideomycetes and Sordariomycetes), 16 orders, 41 families and 60 genera. One order Planisphaeriales, four families Leptosphaerioidaceae, Neoleptosporellaceae, Planisphaeriaceae and Profundisphaeriaceae, ten genera Conicosphaeria, Karstiomyces, Leptosphaerioides, Neoceratosphaeria, Neodiaporthe, Neodictyospora, Planisphaeria, Profundisphaeria, Stellatus and Truncatascus, and 34 species (Amphisphaeria karsti, Anteaglonium hydei, Atractospora terrestris, Conicosphaeria vaginatispora, Corylicola hydei, Diaporthe cylindriformispora, Dictyosporium karsti, Hysterobrevium karsti, Karstiomyces guizhouensis, Leptosphaerioides guizhouensis, Lophiotrema karsti, Murispora hydei, Muyocopron karsti, Neoaquastroma guizhouense, Neoceratosphaeria karsti, Neodiaporthe reniformispora, Neodictyospora karsti, Neoheleiosa guizhouensis, Neoleptosporella fusiformispora, Neoophiobolus filiformisporum, Ophioceras guizhouensis, Ophiosphaerella karsti, Paraeutypella longiasca, Paraeutypella karsti, Patellaria guizhouensis, Planisphaeria karsti, Planisphaeria reniformispora, Poaceascoma herbaceum, Profundisphaeria fusiformispora, Pseudocoleophoma guizhouensis, Pseudopolyplosphaeria guizhouensis, Stellatus guizhouensis, Sulcatispora karsti and Truncatascus microsporus) are introduced as new to science. Moreover, 13 new geographical records for China are also reported, which are Acrocalymma medicaginis, Annulohypoxylon thailandicum, Astrosphaeriella bambusae, Diaporthe novem, Hypoxylon rubiginosum, Ophiosphaerella agrostidis, Ophiosphaerella chiangraiensis, Patellaria atrata, Polyplosphaeria fusca, Psiloglonium macrosporum, Sarimanas shirakamiense, Thyridaria broussonetiae and Tremateia chromolaenae. Additionally, the family Eriomycetaceae was resurrected as a non-lichenized family and accommodated within Monoblastiales. Detailed descriptions and illustrations of all these taxa are provided.

Over the footprints of Italian mycology with emphasis on plant-associated Ascomycota

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May 2023

plant-associated Italian Ascomycota

Microfungi associated with ornamental palms: Byssosphaeria phoenicis sp. nov. (Melanommataceae) and Pseudocoleophoma rhapidis sp. nov. (Dictyosporiaceae) from south China

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Full-text available

Oct 2022

The health and attractiveness of ornamental plants play key role in gardening and horticulture. They affect both the economical and ornamental value of the plants. In this study, we collected and isolated microfungi associated with dead petioles of dwarf date palm and living leaves of lady palm from Zhongkai University Garden, Guangzhou, China. Phylogenetic relationships were investigated based on multi-locus phylogeny of ITS, LSU, SSU and tef-1α sequence data. The morphology of the new collections was compared with all the relevant species. Based on the morphological comparison and multi-locus phylogeny, we introduce Byssosphaeria phoenicis sp. nov. and Pseudocoleophoma rhapidis sp. nov. in Melanommataceae and Dictyosporiaceae, respectively. Detailed descriptions and illustrations are provided with notes discussing allied species in Byssosphaeria and Pseudocoleophoma. Pathogenicity testing of P. rhapidis causing leaf spots on lady palm was proven and Koch’s postulates fulfilled. This is the first phytopathogenic Pseudocoleophoma species. The pathogenicity does not cause dieback, but the leaf spots reduce the aesthetical value of the plants.

New species or records of Cacumisporium, Helicosporium, Monotosporella and Bahusutrabeeja on submerged wood in Hong Kong streams

Article

Full-text available

Mar 2001

Cacumisporium rugosum sp. nov., Helicosporium gigasporum sp. nov. and H. hongkongense sp. nov. are described from submerged wood collected in streams in Hong Kong. Unusual features observed in our collection of Monotosporella rhizoidea are reported, while Bahusutrabeeja dwaya is a new record from streams in Hong Kong.

Ainsworth & Bisby's Dictionary of the Fungi

Article

Full-text available

Jan 1996

The family Pleosporaceae: intergeneric relationships and phylogenetic perspectives based on sequence analyses of partial 28S rDNA

Article

Full-text available

Jul 2006

The Pleosporaceae is an important loculoascomycete family. There has been disagreement, however, regarding the taxonomic placement of many genera within this family. This study investigates phylogenetic relationships among the genera Cochliobolus, Kirschsteiniothelia, Leptosphaerulina, Macroventuria, Pleospora, Pyrenophora, and Wettsteinina. Partial 28S rDNA sequences from taxa within these genera were analyzed with maximum parsimony, likelihood and Bayesian methods. Cochliobolus can be segregated broadly into two groups as previously proposed. Pleospora is polyphyletic in its current sense. Taxa with Stemphylium anamorphs are closely related to Cochliobolus and fit within the Pleosporaceae, whereas the affinities of Pleospora herbarum and P. ambigua are still ambiguous. Pyrenophora constitutes a monophyletic group within the Pleosporaceae, whereas Leptosphaerulina and Macroventuria appear to share phylogenetic affinities with the Leptosphaeriaceae and Phaeosphaeriaceae. Phylogenies indicate that Wettsteinina should be excluded from the Pleosporaceae. Similar findings are reported for Kirschsteiniothelia, which is probably polyphyletic. Anamorphic characters appear to be significant (especially in Cochliobolus) while ascospore morphologies, such as shape and color and substrate occurrence are poor indicators of phylogenetic relationships among these loculoascomycetes.

A class-wide phylogenetic assessment of Dothideomycetes

Article

Full-text available

Jan 2009

Dematiaceous Hyphomycetes

Book

Jan 1971

M.B. Ellis

Freshwater ascomycetes: new or noteworthy species from north temperate lakes in Wisconsin

Article

May 2001

Thirty-one ascomycetes from seven north temperate lakes in Wisconsin are described and illustrated. Among these, one discomycete, Aquadiscula clavispora sp. nov., four pyrenomycetes, Aquadulciospora rhomboidia gen. et sp. nov., Saccardoëlla allequashensis sp. nov., S. horizontalis sp. nov., S. lacustris sp. nov., and one loculoascomycete, Pseudotrichia allequashensis sp. nov., are new. Twelve species are reported from freshwater for the first time.

A Classification of Loculoascomycetes

Article

Sep 1979

Margaret E. Barr

A classification of the Loculoascomycetes is presented in the form of dichotomous keys to 58 families in eight orders. The families Chaeto-thyriaceae, Fenestellaceae, Parmulariaceae, Phaeosphaeriaceae, Pied-raiaceae, Pleomassariaceae, Pyrenophoraceae, Tubeufiaceae, and Venturiaceae are newly described or are validated. For each family, the type genus is designated and representative genera are indicated.

Neotropical Ascomycetes 3. Reinstatement of the Genus Xenolophium and Two New Species from French Guiana

Article

May 1993

S.M. Huhndorf

The genus Xenolophium is recognized for X. pachythele and X. applanatum, both new combinations, and their affinities to Ostropella albocincta are discussed. Two new species X. samuelsii and X. guianense are described from French Guiana. A key to the species of Xenolophium is given.

MAFFT multiple sequence alignment software version 7: improvements in performance and usability

Article

Jan 2013
MOL BIOL EVOL

Standley DM

Fungi Associated with Pandanaceae

Book

Jan 2012

Acknowledgements Dedication 1.The Current Understanding of Fungi Associated with Pandanaceae 2. Methodology 3. Teleomorphic Microfungi Associated with Pandanaceae 4. Anamorphic Microfungi Associated with Pandanaceae 5. List of Fungi Associated with Pandanaceae.