Available via license: CC BY-NC-ND 4.0
Content may be subject to copyright.
Revision of the Massarineae (Pleosporales,Dothideomycetes)
K. Tanaka
1*
, K. Hirayama
2
, H. Yonezawa
1
, G. Sato
1
, A. Toriyabe
1
, H. Kudo
1
, A. Hashimoto
1
,
3
, M. Matsumura
1
, Y. Harada
1
, Y. Kurihara
4
,
T. Shirouzu
5
, and T. Hosoya
5
1
Faculty of Agriculture and Life Science, Hirosaki University, 3 Bunkyo-cho, Hirosaki, Aomori 036-8561, Japan;
2
Apple Experiment Station, Aomori Prefectural Agriculture
and Forestry Research Center, 24 Fukutami, Botandaira, Kuroishi, Aomori 036-0332, Japan;
3
The United Graduate School of Agricultural Sciences, Iwate University, 18-8
Ueda 3 chome, Morioka 020-8550, Japan;
4
OPBIO Factory, 5-8 Suzaki, Uruma, Okinawa 904-2234, Japan;
5
Department of Botany, National Museum of Nature and
Science, 4-1-1 Amakubo, Tsukuba, Ibaraki 305-0005, Japan
*Correspondence: K. Tanaka,
k-tanaka@hirosaki-u.ac.jp
Abstract: We here taxonomically revise the suborder Massarineae (Pleosporales,Dothideomycetes,Ascomycota). Sequences of SSU and LSU nrDNA and the
translation elongation factor 1-alpha gene (tef1) are newly obtained from 106 Massarineae taxa that are phylogenetically analysed along with published sequences of 131
taxa in this suborder retrieved from GenBank. We recognise 12 families and five unknown lineages in the Massarineae. Among the nine families previously known, the
monophyletic status of the Dictyosporiaceae,Didymosphaeriaceae,Latoruaceae,Macrodiplodiopsidaceae,Massarinaceae,Morosphaeriaceae, and Trematosphaer-
iaceae was strongly supported with bootstrap support values above 96 %, while the clades of the Bambusicolaceae and the Lentitheciaceae are moderately supported.
Two new families, Parabambusicolaceae and Sulcatisporaceae, are proposed. The Parabambusicolaceae is erected to accommodate Aquastroma and Parabambusicola
genera nova, as well as two unnamed Monodictys species. The Parabambusicolaceae is characterised by depressed globose to hemispherical ascomata with or without
surrounding stromatic tissue, and multi-septate, clavate to fusiform, hyaline ascospores. The Sulcatisporaceae is established for Magnicamarosporium and Sulcatispora
genera nova and Neobambusicola. The Sulcatisporaceae is characterised by subglobose ascomata with a short ostiolar neck, trabeculate pseudoparaphyses, clavate
asci, broadly fusiform ascospores, and ellipsoid to subglobose conidia with or without striate ornamentation. The genus Periconia and its relatives are segregated from
the Massarinaceae and placed in a resurrected family, the Periconiaceae. We have summarised the morphological and ecological features, and clarified the accepted
members of each family. Ten new genera, 22 new species, and seven new combinations are described and illustrated. The complete ITS sequences of nrDNA are also
provided for all new taxa for use as barcode markers.
Key words: Coelomycetes, Freshwater ascomycetes, Helminthosporium, Holomorph, Hyphomycetes, Massarina,Periconia,Spegazzinia.
Taxonomic novelties: New families: Parabambusicolaceae Kaz. Tanaka & K. Hiray., Sulcatisporaceae Kaz. Tanaka & K. Hiray; New genera: Aquastroma Kaz. Tanaka
& K. Hiray., Clypeoloculus Kaz. Tanaka & K. Hiray., Fuscostagonospora Kaz. Tanaka & K. Hiray., Gregarithecium Kaz. Tanaka & K. Hiray., Magnicamarosporium Kaz.
Tanaka & K. Hiray., Neoophiosphaerella Kaz. Tanaka & K. Hiray., Parabambusicola Kaz. Tanaka & K. Hiray., Pseudocoleophoma Kaz. Tanaka & K. Hiray.,
Pseudoxylomyces Kaz. Tanaka & K. Hiray., Sulcatispora Kaz. Tanaka & K. Hiray; New species: Aquastroma magniostiolata Kaz. Tanaka & K. Hiray., Aquilomyces
rebunensis Kaz. Tanaka & K. Hiray., Clypeoloculus akitaensis Kaz. Tanaka & K. Hiray., Clypeoloculus hirosakiensis Kaz. Tanaka & K. Hiray., Clypeoloculus
microsporus Kaz. Tanaka & K. Hiray., Clypeoloculus towadaensis Kaz. Tanaka & K. Hiray., Dictyosporium pseudomusae Kaz. Tanaka, G. Sato & K. Hiray.,
Fuscostagonospora sasae Kaz. Tanaka & K. Hiray., Gregarithecium curvisporum Kaz. Tanaka & K. Hiray., Helminthosporium massarinum Kaz. Tanaka, K. Hiray. &
Shirouzu, Keissleriella breviasca Kaz. Tanaka & K. Hiray., Keissleriella quadriseptata Kaz. Tanaka & K. Hiray., Keissleriella yonaguniensis Kaz. Tanaka & K. Hiray.,
Lentithecium pseudoclioninum Kaz. Tanaka & K. Hiray., Magnicamarosporium iriomotense Kaz. Tanaka & K. Hiray., Periconia homothallica Kaz. Tanaka & K. Hiray.,
Periconia pseudodigitata Kaz. Tanaka & K. Hiray., Pseudocoleophoma calamagrostidis Kaz. Tanaka & K. Hiray., Pseudocoleophoma polygonicola Kaz. Tanaka & K.
Hiray., Stagonospora pseudoperfecta Kaz. Tanaka & K. Hiray., Sulcatispora acerina Kaz. Tanaka & K. Hiray., Sulcatispora berchemiae Kaz. Tanaka & K.
Hiray; New combinations: Lentithecium clioninum (Kaz. Tanaka, Sat. Hatak. & Y. Harada) Kaz. Tanaka & K. Hiray., Neoophiosphaerella sasicola (Nagas. & Y.
Otani) Kaz. Tanaka & K. Hiray., Parabambusicola bambusina (Teng) Kaz. Tanaka & K. Hiray., Pseudoxylomyces elegans (Goh, W.H. Ho, K.D. Hyde & K.M. Tsui)
Kaz. Tanaka & K. Hiray., Setoseptoria arundinacea (Sowerby) Kaz. Tanaka & K. Hiray., Setoseptoria magniarundinacea (Kaz. Tanaka & Y. Harada) Kaz. Tanaka &
K. Hiray., Stagonospora bicolor (D. Hawksw., W.J. Kaiser & Ndimande) Kaz. Tanaka & K. Hiray; Epitypifications (basionyms): Phaeosphaeria arundinacea var.
brevispora Nagas. & Y. Otani, Phaeosphaeria sasicola Nagas. & Y. Otani.
Available online 18 November 2015; http://dx.doi.org/10.1016/j.simyco.2015.10.002. Hard copy: September 2015.
INTRODUCTION
The Pleosporales is the largest order in the Dothideomycetes.
Two suborders have been recognised, viz. the Pleosporineae
and the Massarineae, containing more than 19 families that
presently still lack subordinal assignment (Hyde et al. 2013). One
suborder, the Pleosporineae, includes many economically
important plant pathogens, Alternaria,Bipolaris,Didymella,
Leptosphaeria,Parastagonospora, and Pyrenophora, for
example, containing huge species diversity (Manamgoda et al.
2011, Zhang et al. 2012, Woudenberg et al. 2013). In this sub-
order, the phylogenetic relationships within each family, the
morphological circumscriptions including those of asexual
morphs, their evolutionary trend as plant pathogens, biogeog-
raphy, and speciation are well established and documented
(Rouxel & Balesdent 2005, Peever 2007, Zhang et al. 2009b,
Ohm et al. 2012, Grandaubert et al. 2014). In contrast, for the
other suborder, the Massarineae, much fundamental information
including the taxonomic framework, phylogenetic relationships,
biology, and species diversity are poorly understood. The Mas-
sarineae was originally established by Barr (1979) to accom-
modate the Massarinaceae and the Arthopyreniaceae, but this
suborder has long been disregarded in the classification of
bitunicate ascomycetes (Hawksworth et al. 1983, 1995, Barr
1987, Eriksson & Winka 1998, Kirk et al. 2008, Lumbsch &
Huhndorf 2010).
Peer review under responsibility of CBS-KNAW Fungal Biodiversity Centre.
Copyright © 2015, CBS-KNAW Fungal Biodiversity Centre. Production and hosting by ELSEVIER B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/
licenses/by-nc-nd/4.0/).
available online at www.studiesinmycology.org STUDIES IN MYCOLOGY 82: 75–136.
75
Studies in Mycology
The family Massarinaceae was established by Munk (1956)
to encompass four genera, Massarina,Metasphaeria,Pseudo-
trichia, and Trichometasphaeria. The thin basal peridium and
dark tissue (= clypeus) around the ostiole found in the ascomata
of these genera were emphasised as familial characters.
Although this proposal was accepted by many researchers
(Eriksson 1981, Boise 1985), the characters emphasised by
Munk (1956) for the family also exist in several other groups of
bitunicate ascomycetes, such as Didymosphaeria (Didymos-
phaeriaceae), Salsuginea (Salsugineaceae) and Roussoella
(Roussoellaceae). The Massarinaceae, therefore, has been
treated as a synonym of the Pleosporaceae (Bose 1961, Luttrell
1973, Von Arx & Müller 1975)orLophiostomataceae (Barr 1987,
1992). Similarly, taxonomic circumscription of the genus Mas-
sarina has also been repeatedly revised (see Notes in Massar-
ina), and as a consequence of a clear taxonomic definition of
Massarina, the family Massarinaceae has been regarded as a
natural lineage. Several related genera, e.g., Aquaticheirospora,
Dictyosporium, and Periconia (Tsui et al. 2006, Kodsueb et al.
2007, Schoch et al. 2009), have been assigned to the Massar-
inaceae as circumscribed by Eriksson & Hawksworth (2003), and
the concept of the family has been revised (Hyde et al. 2013).
In a recent molecular study on the Pleosporales (Zhang et al.
2012), the suborder Massarineae was resurrected as a sister to
the Pleosporineae, and was emended to include five families, the
Lentitheciaceae (Zhang et al. 2009b), Massarinaceae (Munk
1956), Montagnulaceae (Barr 2001; later synonymised under
Didymosphaeriaceae,Ariyawansa et al. 2014), Moro-
sphaeriaceae (Suetrong et al. 2009), and Trematosphaeriaceae
(Suetrong et al. 2011b). More recently, the Bambusicolaceae
(Hyde et al. 2013), Dictyosporiaceae (nom. prov., see Liu et al.
2015), Latoruaceae and Macrodiplodiopsidaceae (Crous et al.
2015a) have been added to the suborder. Most of these fam-
ilies have been recognised from the results of recent molecular
studies. Only a few members of each family are currently known,
and thus the morphological characteristics and phylogenetic
relationships within each family are not fully understood.
Consequently, several new genera belonging to the Massar-
ineae, such as Ascorhombispora (Cai & Hyde 2007b), Inflatis-
pora (Zhang et al. 2011), and Noosia (Crous et al. 2011a), have
been published but their phylogenetic placements at familial level
remain obscure (Zhang et al. 2012, Hyde et al. 2013).
Our aims were to reveal the species diversity within the
Massarineae and to establish a taxonomic framework within this
suborder for understanding the relationships among the Mas-
sarineae. To this end we examined 106 specimens/isolates
belonging to this suborder, and analysed their morphology and
partial DNA sequences of the small and large subunit nuclear
ribosomal DNA (SSU and LSU nrDNA) and the translation
elongation factor 1-alpha gene (tef1).
MATERIALS AND METHODS
Morphological studies
Leaf and twig specimens were collected from various plants in
Japan, and deposited in the herbarium of Hirosaki University
(HHUF). Measurements of all structures except for ascomata/
conidiomata were taken from material mounted in distilled water.
India ink or Black-Blue ink in distilled water was added to water
mounts to detect gelatinous sheaths or appendages around
spores. To observe the internal structure of strongly melanised
spores, 5 % sodium hypochlorite solution (NaClO) was used for
the bleaching of spores as described in Eriksson (1989). The
position of the primary septum of spores was noted using the
decimal system (Shoemaker 1984), and the numbers of spore
septa were recorded as “septa of upper hemisphere + the pri-
mary septum + septa of lower hemisphere”. To observe sporo-
carp structure, ascomata/conidiomata were boiled in water for a
few minutes, sectioned using a freezing microtome (HM 400R;
MICROM, Germany), and mounted in diluted lactophenol cotton
blue. Morphology was observed using differential interference
and phase contrast microscopy (Olympus BX53, Japan).
Single spore cultures were obtained following the methods of
Tubaki (1978). Ninety-nine cultures on 2 % potato-dextrose agar
(PDA) or potato-carrot agar (PCA) were prepared from the col-
lections and were deposited in the Japan Collection of Micro-
organisms (JCM), the National Institute of Agrobiological
Sciences, Japan (MAFF), and the CBS-KNAW Fungal Biodi-
versity Centre (Centraalbureau voor Schimmelcultures; CBS).
An additional seven strains were obtained from the CBS culture
collection (Table 1). Growth rate and colony characteristics were
recorded from cultures grown on PDA within 2 or 4 wk at 20 °C in
the dark. Colours were designated according to Rayner (1970).
Induction of sexual/asexual sporulation was attempted by
culturing isolates on rice straw agar (RSA; Tanaka & Harada
2003a) and/or incubating small pieces of colony in sterilised
water (Scheuer 1991). Nomenclatural novelties were deposited
in MycoBank (Crous et al. 2004).
Molecular phylogenetic analysis
A total of 106 isolates were used for DNA extraction (Table 1).
DNA from mycelia was extracted using the ISOPLANT Kit
(Nippon Gene, Tokyo, Japan) following the manufacturer’s in-
structions. Partial SSU and LSU nrDNA, and tef1 were
sequenced to elucidate phylogenetic relationships of the isolates
for considering familial and generic classifications. The complete
internally transcribed spacer (ITS) regions of nrDNA were also
obtained for use as DNA barcode markers (Schoch et al. 2012),
although the sequences were not used for phylogenetic recon-
struction (Table 1). Four primer sets, NS1–NS4 (White et al.
1990), LR0R–LR7 (Rehner & Samuels 1994), EF1-983F–
EF1-2218R (Rehner & Buckley 2005), and ITS1–ITS4 (White
et al. 1990) were used for the amplification of SSU, LSU, tef1,
and ITS, respectively. DNA fragments were amplified and
sequenced following the methods described by Tanaka et al.
(2009). Newly obtained sequences have been deposited in
GenBank (Table 1). These sequences together with those
retrieved from GenBank (Table 2) were aligned by MUSCLE
included in the program Molecular Evolutionary Genetic Analysis
(MEGA) v. 6 (Tamura et al. 2013) and manually adjusted to
optimise the alignment. Hysterobrevium mori and Hysterium
pulicare, both belonging to the Hysteriales, were designated as
outgroup taxa. The alignments used were deposited in Tree-
BASE (http://www.treebase.org). Phylogenetic analyses were
conducted based on maximum likelihood (ML) method. The
optimum substitution models for each dataset were estimated by
Kakusan4 (Tanabe 2011), based on the Akaike information cri-
terion (AIC; Akaike 1974) for ML analyses. The ML analyses
were performed with TreeFinder Mar 2011 (Jobb 2011) based on
TANAKA ET AL.
76
Table 1. Cultures and GenBank accession numbers of Massarineae obtained in this study.
Taxon Family
1
Original no. Culture no. Specimen no.
2
GenBank accession no.
3
Notes
4
SSU LSU tef1 ITS
Aquastroma magniostiolata Par KT 2485 CBS 139680 = JCM 19429 = MAFF 243824 HHUF 30122
HT
AB797220 AB807510 AB808486 LC014540 A
Aquilomyces rebunensis Mor KT 732-2 CBS 139684 = JCM 19427 = MAFF 243862 HHUF 27556
HT
AB797252 AB807542 AB808518 AB809630 A
Bactrodesmium cubense IS –CBS 680.96 = JCM 14126 –AB797218 AB807508 AB808484 LC014541 C
Clypeoloculus akitaensis Mor KT 788 CBS 139681 = JCM 19424 = MAFF 239467 HHUF 27557
HT
AB797253 AB807543 AB808519 AB809631 A
C. hirosakiensis Mor KT 1283 CBS 139682 = JCM 19425 = MAFF 243864 HHUF 30144
HT
AB797260 AB807550 AB808526 AB809638 A
C. microsporus Mor KT 1131 CBS 139683 = JCM 19426 = MAFF 243863 HHUF 30143
HT
AB797245 AB807535 AB808510 AB811451 A
C. towadaensis Mor KT 1340 CBS 139685 = JCM 19428 = MAFF 243865 HHUF 30145
HT
AB797259 AB807549 AB808525 AB809637 A
Dictyosporium aff. bulbosum Dic KH 375 JCM 19403 = MAFF 243829 HHUF 30127 AB797224 AB807514 AB808490 LC014542 C
D. aff. heptasporum Dic KH 332 JCM 19406 = MAFF 243828 HHUF 30126 AB797223 AB807513 AB808489 LC014543 C
D. bulbosum Dic yone 221 MAFF 243835 HHUF 29990 AB797221 AB807511 AB808487 LC014544 C
D. digitatum Dic KH 401 JCM 19404 = MAFF 243830 HHUF 30128 AB797225 AB807515 AB808491 LC014545 C
D. digitatum Dic KT 2660 JCM 19405 = MAFF 243833 HHUF 30131 AB797228 AB807518 AB808494 LC014546 C
D. digitatum Dic yone 280 MAFF 243837 HHUF 30093 AB797222 AB807512 AB808488 LC014547 C
D. hughesii Dic KT 1847 JCM 19407 = MAFF 243832 HHUF 30130 AB797227 AB807517 AB808493 LC014548 C
D. pseudomusae Dic KH 412 JCM 19408 = MAFF 243831 HHUF 30129
PT
AB797226 AB807516 AB808492 LC014549 C
D. pseudomusae Dic yone 234 CBS 139686 = JCM 19409 = MAFF 243836 HHUF 30133
HT
AB797230 AB807520 AB808496 LC014550 C
D. tetrasporum Dic KT 2865 JCM 19410 = MAFF 243834 HHUF 30132 AB797229 AB807519 AB808495 LC014551 C
Fuscostagonospora sasae IS KT 1467 CBS 139687 = JCM 13104 = MAFF 239614 HHUF 29106
HT
AB797258 AB807548 AB808524 AB809636 A
Gregarithecium curvisporum Dic KT 922 CBS 139688 = JCM 19411 = MAFF 243838 HHUF 30134
HT
AB797257 AB807547 AB808523 AB809644 A
Helicascus aquaticus Mor KT 1544 JCM 19423 = MAFF 243866 HHUF 30146 AB797242 AB807532 AB808507 AB809627 A
H. elaterascus Mor KT 2673 MAFF 243867 HHUF 30147 AB797243 AB807533 AB808508 AB809626 A
H. elaterascus Mor KT 2682 CBS 139689 HHUF 30451 LC014603 LC014608 LC014613 LC014552 A
H. thalassioideus Mor –CBS 110441 = JCM 14147 –AB797267 AB807557 AB808533 LC014553 A
H. thalassioideus Mor KH 242 JCM 17526 = NBRC 107811 HHUF 30069 AB797268 AB807558 AB808534 LC014554 A
Helminthosporium dalbergiae Mas H 4628 (= TS 36) MAFF 243853 HHUF 27971 AB797231 AB807521 AB808497 LC014555 C
H. magnisporum Mas H 4627 (= TS 33) MAFF 239278 HHUF 27968
HT
AB797232 AB807522 AB808498 AB811452 C
H. massarinum Mas KT 838 JCM 13094 = MAFF 239604 HHUF 27573
PT
AB797233 AB807523 AB808499 AB809628 A
H. massarinum Mas KT 1564 CBS 139690 = JCM 13095 = MAFF 239605 HHUF 29089
HT
AB797234 AB807524 AB808500 AB809629 A
Helminthosporium sp. Mas H 4743 (= TS 68) MAFF 243856 HHUF 28248 AB797236 AB807526 ––C
Helminthosporium sp. Mas yone 38 MAFF 243857 HHUF 29740 AB797237 AB807527 AB808502 –C
Helminthosporium sp. Mas yone 63 MAFF 243858 HHUF 29741 AB797238 AB807528 AB808503 –C
(continued on next page)
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 77
Table 1. (Continued).
Taxon Family
1
Original no. Culture no. Specimen no.
2
GenBank accession no.
3
Notes
4
SSU LSU tef1 ITS
H. velutinum Mas H 4626 (= TS 28) MAFF 243854 HHUF 27966 AB797240 AB807530 AB808505 LC014556 C
H. velutinum Mas H 4739 (= TS 58) MAFF 243855 HHUF 28243 AB797235 AB807525 AB808501 LC014557 C
H. velutinum Mas yone 96 MAFF 243859 HHUF 30140 AB797239 AB807529 AB808504 LC014558 C
Karstenula rhodostoma Did –CBS 691.94 UPS (F-141152) 425947 AB797241 AB807531 AB808506 LC014559 A
Katumotoa bambusicola Len KT 1517a JCM 13131 = MAFF 239641 HHUF 28661
PT
AB524454
1
AB524595
1
AB539108
2
LC014560 A
Keissleriella breviasca Len KT 540 JCM 19413 = MAFF 239476 HHUF 27715
PT
AB797296 AB807586 AB808565 AB811453 A
K. breviasca Len KT 581 JCM 19414 = MAFF 243843 HHUF 27717
PT
AB797297 AB807587 AB808566 AB811454 A
K. breviasca Len KT 649 CBS 139691 = JCM 19415 = MAFF 243844 HHUF 27718
HT
AB797298 AB807588 AB808567 AB811455 A
K. culmifida Len KT 2308 JCM 19416 = MAFF 243848 HHUF 30135 AB797301 AB807591 AB808570 LC014561 A
K. culmifida Len KT 2642 JCM 19417 = MAFF 243849 HHUF 30136 AB797302 AB807592 AB808571 LC014562 A
K. gloeospora Len KT 829 MAFF 239474 HHUF 27704 AB797299 AB807589 AB808568 LC014563 A
K. quadriseptata Len KT 2292 CBS 139692 = JCM 19418 = MAFF 243850 HHUF 30137
HT
AB797303 AB807593 AB808572 AB811456 A
Keissleriella sp. Len KT 895 JCM 19420 = MAFF 243845 HHUF 27705 AB797300 AB807590 AB808569 –A
K. taminensis Len KT 571 MAFF 243846 HHUF 27707 AB797305 AB807595 AB808574 LC014564 A
K. taminensis Len KT 594 MAFF 243847 HHUF 27709 AB797306 AB807596 ––A
K. taminensis Len KT 678 MAFF 239475 HHUF 27711 AB797307 AB807597 AB808575 LC014565 A
K. yonaguniensis Len KT 2604 CBS 139693 = JCM 19419 = MAFF 243851 HHUF 30138
HT
AB797304 AB807594 AB808573 AB811457 A
Lentithecium clioninum Len KT 1149A CBS 139694 = JCM 12703 = MAFF 239293 HHUF 28199
HT
AB797250 AB807540 AB808515 LC014566 A
L. clioninum Len KT 1220 MAFF 243839 HHUF 28213
PT
AB797251 AB807541 AB808516 LC014567 A
L. pseudoclioninum Len KT 1111 JCM 19421 = MAFF 243840 HHUF 29053
PT
AB797254 AB807544 AB808520 AB809632 A
L. pseudoclioninum Len KT 1113 CBS 139695 = JCM 19422 = MAFF 243841 HHUF 29055
HT
AB797255 AB807545 AB808521 AB809633 A
Magnicamarosporium iriomotense Sul KT 2822 CBS 139696 = JCM 19402 = MAFF 243827 HHUF 30125
HT
AB797219 AB807509 AB808485 AB809640 C
Massarina cisti Mas –CBS 266.62 = JCM 14140 ZT (Hütter & Loeffler)
HT
AB797249 AB807539 AB808514 LC014568 A
M. eburnea Mas H 3953 CBS 139697 = JCM 14422 HHUF 26621 AB521718
3
AB521735
3
AB808517 LC014569 A
Monodictys capensis IS HR 1 CBS 134928 = VKM F-4506 HHUF 29712 AB797261 AB807551 AB808527 LC014570 C
Monodictys sp. Par JO 10 MAFF 243825 HHUF 30123 AB797262 AB807552 AB808528 –C
Monodictys sp. Par KH 331 MAFF 243826 HHUF 30124 AB797263 AB807553 AB808529 –C
Morosphaeria ramunculicola Mor KH 220 NBRC 107813 HHUF 30070 AB797264 AB807554 AB808530 –A
M. velatispora Mor KH 218 JCM 17529 = NBRC 107814 HHUF 30072 AB797265 AB807555 AB808531 LC014571 A
M. velatispora Mor KH 221 JCM 17530 = NBRC 107812 HHUF 30073 AB797266 AB807556 AB808532 LC014572 A
Neokalmusia brevispora Did KT 1466 CBS 120248 = JCM 13543 = MAFF 239276 HHUF 28229 AB524459
1
AB524600
1
AB539112
1
LC014573 A
N. brevispora Did KT 2313 NBRC 106240 HHUF 30016
ET
AB524460
1
AB524601
1
AB539113
2
LC014574 A
N. scabrispora Did KT 1023 CBS 120246 = JCM 12851 = MAFF 239517 HHUF 28608 AB524452
1
AB524593
1
AB539106
2
LC014575 A
TANAKA ET AL.
78
Table 1. (Continued).
Taxon Family
1
Original no. Culture no. Specimen no.
2
GenBank accession no.
3
Notes
4
SSU LSU tef1 ITS
N. scabrispora Did KT 2202 NBRC 106237 HHUF 30013 AB524453
1
AB524594
1
AB539107
2
LC014576 A
Neoophiosphaerella sasicola Len KT 1706 CBS 120247 = JCM 13134 = MAFF 239644 HHUF 29443
ET
AB524458
1
AB524599
1
AB539111
2
LC014577 A
Parabambusicola bambusina Par H 4321 MAFF 239462 HHUF 26590 AB797246 AB807536 AB808511 LC014578 A
P. bambusina Par KH 139 MAFF 243823 HHUF 30121 AB797247 AB807537 AB808512 LC014579 A
P. bambusina Par KT 2637 MAFF 243822 HHUF 30120 AB797248 AB807538 AB808513 LC014580 A
Paraphaeosphaeria michotii Did KT 2222 MAFF 243861 HHUF 30142 AB797269 AB807559 AB808535 AB809639 A
Periconia byssoides Per H 4600 (= TS 29) MAFF 243872 HHUF 28238 AB797280 AB807570 AB808546 LC014581 C
P. byssoides Per H 4432 MAFF 243869 –AB797279 AB807569 AB808545 LC014582 C
P. byssoides Per H 4853 (= TS 60) MAFF 243873 –AB797281 AB807571 AB808547 LC014583 C
P. digitata Per –CBS 510.77 –AB797271 AB807561 AB808537 LC014584 C
P. homothallica Per KT 916 CBS 139698 = JCM 13100 = MAFF 239610 HHUF 29105
HT
AB797275 AB807565 AB808541 AB809645 A
P. igniaria Per –CBS 379.86 –AB797276 AB807566 AB808542 LC014585 C
P. igniaria Per –CBS 845.96 = JCM 14142 –AB797277 AB807567 AB808543 LC014586 C
P. pseudobyssoides Per H 4151 MAFF 243868 –AB797278 AB807568 AB808544 LC014587 C
P. pseudobyssoides Per H 4790 ( = TS 102) MAFF 243874 HHUF 28257 AB797270 AB807560 AB808536 LC014588 C
P. pseudodigitata Per KT 644 JCM 13164 = MAFF 239674 HHUF 27569
PT
AB797272 AB807562 AB808538 LC014589 A
P. pseudodigitata Per KT 1195A JCM 13165 = MAFF 239675 HHUF 29368
PT
AB797273 AB807563 AB808539 LC014590 A
P. pseudodigitata Per KT 1395 CBS 139699 = JCM 13166 = MAFF 239676 HHUF 29370
HT
AB797274 AB807564 AB808540 LC014591 A
Periconia sp. Per KT 1820A MAFF 243870 HHUF 30148 AB797282 AB807572 AB808548 –C
Periconia sp. Per KT 1825 MAFF 243871 HHUF 30149 AB797283 AB807573 AB808549 –C
Pseudocoleophoma calamagrostidis Dic KT 3284 CBS 139700 HHUF 30450
HT
LC014604 LC014609 LC014614 LC014592 A
P. polygonicola Dic KT 731 CBS 139701 = JCM 19412 = MAFF 239468 HHUF 27558
HT
AB797256 AB807546 AB808522 AB809634 A
Setoseptoria arundinacea Len KT 552 MAFF 239460 HHUF 27543 AB797284 AB807574 AB808550 LC014594 A
S. arundinacea Len KT 600 MAFF 243842 HHUF 27544 AB797285 AB807575 AB808551 LC014595 A
S. magniarundinacea Len KT 1174 CBS 139702 = MAFF 239294 HHUF 28293
HT
AB797286 AB807576 AB808552 LC014596 A
Spegazzinia deightonii Did yone 66 MAFF 243876 HHUF 30150 AB797291 AB807581 AB808557 –C
S. deightonii Did yone 212 MAFF 243877 HHUF 30151 AB797292 AB807582 AB808558 –C
Spegazzinia sp. Did yone 279 MAFF 243878 HHUF 30152 AB797293 AB807583 AB808559 –C
S. tessarthra Did SH 287 MAFF 243875 HHUF 27691 AB797294 AB807584 AB808560 –C
Stagonospora perfecta Mas KT 1726A JCM 13099 = MAFF 239609 HHUF 29095 AB797289 AB807579 AB808555 AB809642 A
S. pseudoperfecta Mas KT 889 CBS 120236 = JCM 13097 = MAFF 239607 HHUF 29087
HT
AB797287 AB807577 AB808553 AB809641 A
Stagonospora sp. Mas KT 903 CBS 120237 = JCM 13098 = MAFF 239608 HHUF 29088 AB797288 AB807578 AB808554 –A
(continued on next page)
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 79
the models selected by AIC (separate model among genes and
proportional model among codons, HKY85+G for SSU, J2+G for
LSU, GTR+G for the first codon of tef1, TVM+G for the second
codon of tef1, and GTR+G for the third codon of tef1). Bootstrap
proportions (BP) were obtained by 1000 bootstrap replications.
RESULTS
Molecular phylogenetic analysis
Approximately 940–1 750 bp of SSU, 870–1 330 bp of LSU
nrDNA, 830–940 bp of tef1, and 500–900 bp of ITS sequences
were determined for 106 isolates of fungi within the Massarineae.
Analyses of different gene datasets were performed individually,
but no topological conflict was observed at familial level with the
exception of Lentitheciaceae, which was polyphyletic in the tef1
tree (data not shown). A combined dataset of SSU, LSU, and tef1
sequences was generated after excluding insertions of several
species which corresponded to positions 493–1 005 of Mono-
dictys capensis (GenBank AB797261) and positions
1 286–1 651 of Magnicamarosporium iriomotense (GenBank
AB797219) in the SSU, and positions 836–892 of Montagnula
spartii (GenBank GU205225) and positions 871–924 of Hys-
terium pulicare (GenBank FJ161201) in the LSU sequences. The
combined dataset consisted of 243 taxa and 3 386 characters, of
which 28 % were missing and gap characters. The alignment
had 82 % representation for SSU, 100 % for LSU and 60 % for
tef1. BP support of each familial clade in the LSU tree was
generally improved by adding the SSU and tef1 dataset with
missing data, e.g., from 82 % to 100 % in the Massarinaceae,
and from 90 % to 100 % in the Trematosphaeriaceae.
The ML tree of Massarineae based on the SSU and LSU
nrDNA and tef1 regions with the highest log likelihood
(−36965.05250) is shown in Fig. 1. A total of 237 taxa of the
Massarineae formed a clade (with 98 % BP support) and were
scattered in 12 familial clades and five unknown clades. Seven
families previously recognised, namely the Dictyosporiaceae
(nom. prov., 100 % BP), Didymosphaeriaceae (99 % BP),
Latoruaceae (99 % BP), Macrodiplodiopsidaceae (96 % BP),
Massarinaceae (100 % BP), Morosphaeriaceae (98 % BP), and
Trematosphaeriaceae (100 % BP) were highly supported as
independent monophyletic groups. The Bambusicolaceae and
Lentitheciaceae received moderate BP support, 87 % and 71 %,
respectively. We erected two new families, the Para-
bambusicolaceae (81 % BP) and Sulcatisporaceae (97 % BP), to
accommodate several genera, which cannot be placed in any of
the existing families in the Massarineae. Although species in the
genus Periconia have been treated as members of the Mas-
sarinaceae (Zhang et al. 2012, Hyde et al. 2013), we have placed
them in a distinct family, the Periconiaceae (100 % BP), which
was established by Nannizzi (1934).
Taxonomy
As a result of morphological comparisons and phylogenetic
analyses of 106 strains, along with sequences from 131 taxa
obtained from GenBank, at least 12 families including two new
families (the Parabambusicolaceae and Sulcatisporaceae) are
recognised. Ten new genera, 22 new species, and seven new
combinations are proposed. Taxa are arranged in alphabetical
Table 1. (Continued).
Taxon Family
1
Original no. Culture no. Specimen no.
2
GenBank accession no.
3
Notes
4
SSU LSU tef1 ITS
S. tainanensis Mas KT 1866 MAFF 243860 HHUF 30141 AB797290 AB807580 AB808556 AB809643 A
Sulcatispora acerina Sul KT 2982 CBS 139703 HHUF 30449
HT
LC014605 LC014610 LC014615 LC014597 A
S. berchemiae Sul KT 1607 CBS 139704 = JCM 13101 = MAFF 239611 HHUF 29097
HT
AB797244 AB807534 AB808509 AB809635 A
Tingoldiago graminicola Len KH 68 JCM 16485 = NBRC 106131 HHUF 30009
HT
AB521726
3
AB521743
3
AB808561 LC014598 A
T. graminicola Len KH 155 JCM 16486 = NBRC 106132 HHUF 30010
PT
AB521728
3
AB521745
3
AB808562 LC014599 A
T. graminicola Len KT 891 MAFF 239472 HHUF 27882
PT
AB521727
3
AB521744
3
AB808563 LC014600 A
Trematosphaeria pertusa Tre KT 1496 JCM 19430 = MAFF 243879 HHUF 30153 AB797295 AB807585 AB808564 AB809646 A
T. pertusa Tre KT 3314 CBS 139705 HHUF 30452 LC014606 LC014611 LC014616 LC014601 A
T. pertusa Tre KT 3315 CBS 139706 HHUF 30453 LC014607 LC014612 LC014617 LC014602 A
Pseudoxylomyces elegans IS KT 2887 MAFF 243852 HHUF 30139 AB797308 AB807598 AB808576 LC014593 C
1
Abbreviation of family names: Bam (Bambusicolaceae), Dic (Dictyosporaceae), Did (Didymosphaeriaceae), IS (insertae sedis), Len (Lentitheciaceae), Mas (Massarinaceae), Mor (Morosphaeriaceae), Par (Paramorosphaeriaceae), Per (Peri-
coniaceae), Sul (Sulcatisporaceae), and Tre (Trematosphaeriaceae).
2
Specimen with HT (holotype), PT (paratype), and ET (epitype).
3
Sequences obtained in this study are shown in bold. Sequences with
1
from Tanaka et al. (2009),
2
from Schoch et al. (2009), and
3
from Hirayama et al. (2010).
4
Origin of isolates: A, single ascospore; C, single conidium.
TANAKA ET AL.
80
Table 2. Cultures and GenBank accession numbers of Massarineae used for phylogenetic analysis.
Taxon Family
1
Culture no.
2
GenBank accession no.
3
SSU LSU tef1
Alloconiothyrium aptrootii Did CBS 980.95
HT
NS JX496234 NS
Aquaticheirospora lignicola Dic RK-2006a
HT
AY736377 AY736378 NS
Aquilomyces patris Mor CBS 135661
HT
KP184077 KP184041 NS
Asteromassaria pulchra IS CBS 124082 GU296137 GU301800 GU349066
Bambusicola bambusae Bam MFLUCC 11-0614
HT
JX442039 JX442035 NS
B. irregulispora Bam MFLUCC 11-0437
HT
JX442040 JX442036 NS
B. loculata Bam MFLUCC 13-0856
HT
KP761735 KP761729 KP761724
B. massarinia Bam MFLUCC 11-0389
HT
JX442041 JX442037 NS
B. splendida Bam MFLUCC 11-0439
HT
JX442042 JX442038 NS
Bambusistroma didymosporum Per MFLUCC 13-0862
HT
NS KP761730 KP761727
Bimuria novae-zelandiae Did CBS 107.79
HT
AY016338 AY016356 DQ471087
Byssothecium circinans Mas CBS 675.92 GU205235 AY016357 GU349061
Camarographium koreanum Mac CBS 117159
HT
NS JQ044451 NS
“Corynespora”leucadendri Mas CBS 135133
HT
NS KF251654 NS
“C.”olivacea Mas CBS 114450 NS GU301809 GU349014
Cucurbidothis pityophila Did CBS 149.32 U42480 DQ384102 NS
Darksidea alpha Len CBS 135650
HT
KP184049 KP184019 KP184166
D. beta Len CBS 135637
HT
KP184074 KP184023 KP184189
D. gamma Len CBS 135634
HT
KP184073 KP184028 KP184188
D. delta Len CBS 135638
HT
KP184069 KP184024 KP184184
D. epsilon Len CBS 135658
HT
KP184070 KP184029 KP184186
D. zeta Len CBS 135640
HT
KP184071 KP184013 KP184191
Dendryphiella vinosa Dic –EU848589 EU848590 NS
Deniquelata barringtoniae Did MFLUCC 110422
HT
JX254656 JX254655 NS
Dictyosporium alatum Dic ATCC 34953
HT
DQ018080 DQ018101 NS
D. elegans Dic NBRC 32502 DQ018079 DQ018100 NS
D. inflatum Dic NTOU 3855 JQ267361 JQ267363 NS
D. meiosporum Dic MFLUCC 10-0131
HT
KP710946 KP710945 NS
D. stellatum Dic CCFC 241241
HT
NS JF951177 NS
D. strelitziae Dic CBS 123359
HT
NS FJ839653 NS
D. thailandicum Dic MFLUCC 13-0773
HT
NS KP716707 NS
D. toruloides Dic CBS 209.65 DQ018081 DQ018104 NS
Didymocrea sadasivanii Did CBS 438.65
IT
DQ384066 DQ384103 NS
Didymosphaeria rubi-ulmifolii Did MFLUCC 14-0023
HT
KJ436588 KJ436586 NS
“D.”spartii Mas CBS 183.58 GU205250 GU205225 NS
Digitodesmium bambusicola Dic CBS 110279
HT
NS DQ018103 NS
“Diplococcium”asperum Dic CBS 139.95 EF204511 EF204493 NS
Falciformispora lignatilis Tre BCC 21117 GU371834 GU371826 GU371819
F. senegalensis Tre CBS 196.79
HT
KF015636 KF015631 KF015687
F. tompkinsii Tre CBS 200.79
HT
KF015639 KF015625 KF015685
Flavomyces fulophazii Per CBS 135761
HT
KP184082 KP184040 NS
Halomassarina thalassiae Tre BCC 17054 GQ925842 GQ925849 NS
H. thalassiae Tre JK 5262D NS GU301816 GU349011
Helicascus aegyptiacus Mor FWCC 99
HT
KC894852 KC894853 NS
H. aquaticus Mor MFLUCC 10-0918
HT
KC886638 KC886640 NS
H. elaterascus Mor A22-5A = HKUCC 7769 AF053727 AY787934 NS
H. nypae Mor BCC 36751 GU479754 GU479788 GU479854
H. nypae Mor BCC 36752 GU479755 GU479789 GU479855
H. thalassioideus Mor MFLUCC10-0911 KC886637 KC886636 NS
(continued on next page)
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 81
Table 2. (Continued).
Taxon Family
1
Culture no.
2
GenBank accession no.
3
SSU LSU tef1
H. unilocularis Mor MJF14020
HT
NS KP637166 NS
Hysterium pulicare OG CBS 12337 FJ161161 FJ161201 FJ161109
Hysterobrevium mori OG CBS 12356 FJ161155 FJ161196 FJ161104
Inflatispora pseudostromatica IS CBS 123110
HT
JN231132 JN231131 NS
“Kalmusia (Montagnula)”anthostomoides Did CBS 615.86 GU205246 GU205223 NS
K. ebuli Did CBS 123120
NT
JN851818 JN644073 NS
K. italica Did MFLUCC 13–0066
HT
KP325442 KP325441 NS
K. spartii Did MFLUCC 14–0560
HT
KP753953 KP744487 NS
K. variispora Did CBS 121517
HT
NS JX496143 NS
Karstenula rhodostoma Did CBS 690.94 GU296154 GU301821 GU349067
Keissleriella cladophila Len CBS 104.55 GU296155 GU301822 GU349043
K. genistae Len CBS 113798 GU205242 GU205222 NS
K. linearis Len IFRD 2008 FJ795478 FJ795435 NS
K. poagena Len CBS 136767
HT
NS KJ869170 NS
K. sparticola Len MFLUCC 14 –0196
HT
NS KP639571 NS
K. trichophoricola Len CBS 136770
HT
NS KJ869171 NS
Latorua caligans Lat CBS 576.65
HT
NS KR873266 NS
L. grootfonteinensis Lat CBS 369.72
HT
NS KR873267 NS
Lentithecium aquaticum Len CBS 123099
HT
GU296156 GU301823 GU349068
L. fluviatile Len CBS 122367 GU296158 GU301825 GU349074
Leptosphaeria doliolum Lep CBS 505.75 GU296159 GU301827 GU349069
Letendraea eurotioides Did CBS 212.31 NS AY787935 NS
L. helminthicola Did CBS 884.85 AY016345 AY016362 NS
L. padouk Did CBS 485.70 GU296162 AY849951 NS
Lophiostoma arundinis Lop JCM 13550 AB618679 AB618998 LC001737
L. macrostomum Lop JCM 13544 AB618691 AB619010 LC001751
Macrodiplodiopsis desmazieri Mac CBS 140062
ET
NS KR873272 NS
M. desmazieri Mac CBS 221.37 DQ678013 DQ678065 DQ677908
Massarina eburnea Mas CBS 473.64 GU296170 GU301840 GU349040
Montagnula aloes Did CBS 132531
HT
NS JX069847 NS
M. graminicola Did MFLUCC 13–0352
HT
KM658316 KM658315 NS
M. opulenta Did CBS 168.34 AF164370 DQ678086 NS
Morosphaeria ramunculicola Mor BCC 18404 GQ925838 GQ925853 NS
M. velatispora Mor BCC 17059 GQ925841 GQ925852 NS
Multiseptospora thailandica Par MFLUCC 11–0183
HT
KP753955 KP744490 NS
Munkovalsaria appendiculata Did CBS 109027
HT
NS AY772016 NS
Murilentithecium clematidis Len MFLUCC 14-0561
HT
KM408761 KM408759 KM454445
Neobambusicola strelitziae Sul CBS 138869
HT
NS KP004495 NS
Noosia banksiae Per CBS 129526
HT
NS JF951167 NS
Palmiascoma gregariascomum Bam MFLUCC 11–0175
HT
KP753958 KP744495 NS
Paracamarosporium leucandendri Did CBS 123027
HT
NS EU552106 NS
P. psoraleae Did CBS 136628
HT
NS KF777199 NS
Paraconiothyrium estuarinum Did CBS 109850
HT
NS JX496129 NS
“P.”flavescens Dic CBS 178.93 GU238216 GU238075 NS
“P.”fuckelii Did CBS 797.95 GU238204 GU237960 NS
P. maculicutis Did CBS 101461
HT
EU754101 EU754200 NS
P. thysanolaenae Did MFLUCC 10–0550
HT
KP753959 KP744496 NS
“P.”tiliae Did CBS 265.94
LT
EU754040 EU754139 NS
Paraphaeosphaeria michotii Did CBS 652.86 GQ387520 GQ387581 GU456266
TANAKA ET AL.
82
order by family, genus, and species. For the known species a
brief description and/or taxonomic notes are provided.
Dictyosporiaceae nom. prov. (see Liu et al. 2015)
Type genus:Dictyosporium Corda.
Dictyosporium Corda, Weitenweber's Beitr. Nat.: 87. 1836.
Type species:Dictyosporium elegans Corda.
Notes:Dictyosporium species have been reported worldwide
from dead wood and decaying leaves in terrestrial and
Table 2. (Continued).
Taxon Family
1
Culture no.
2
GenBank accession no.
3
SSU LSU tef1
“P.”spartii Did MFLUCC 13–0398
HT
KP711367 KP711362 NS
Periconia macrospinosa Per CBS 135663 KP184080 KP184038 NS
Phaeodothis winteri Did CBS 182.58 GU296183 GU301857 DQ677917
Phragmocamarosporium hederae Len MFLUCC 13-0552
HT
KP842918 KP842915 NS
P. platani Len MFLUCC 14-1191
HT
KP842919 KP842916 NS
“Pithomyces”valparadisiacus Mor CBS 113339 NS EU552152 NS
Pleospora herbarum Ple CBS 191.86 DQ247812 DQ247804 DQ471090
Pleurophoma pleurospora Len CBS 130329
LT
NS JF740327 NS
Poaceascoma helicoides Len MFLUCC 11-0136
HT
KP998463 KP998462 KP998461
Polyschema congolensis Lat CBS 542.73
HT
NS EF204502 NS
P. larviformis Lat CBS 463.88 NS EF204503 NS
P. terricola Lat CBS 301.65
HT
EF204519 EF204504 NS
Pseudocamarosporium brabeji Did CBS 119219
HT
NS EU552104 NS
P. cotinae Did MFLUCC 14–0624
HT
KP753964 KP744505 NS
P. propinquum Did MFLUCC 13-0544
ET
KJ819949 KJ813280 NS
P. tilicola Did MFLUCC 13-0550
HT
KJ819950 KJ813281 NS
Pseudochaetosphaeronema larense Mac CBS 640.73
HT
KF015652 KF015611 KF015684
Pseudodictyosporium elegans Dic CBS 688.93
HT
DQ018084 DQ018106 NS
P. wauense Dic NBRC 30078 DQ018083 DQ018105 NS
Setoseptoria arundinacea Len CBS 619.86 GU296157 GU301824 NS
S. arundinacea Len CBS 123131 GU456298 GU456320 GU456281
S. phragmitis Len CBS 114802
HT
NS KF251752 NS
S. phragmitis Len CBS 114966
PT
NS KF251753 NS
Sporidesmiella fusiformis Did HKUCC 10831 NS DQ408577 NS
“Sporidesmium”tengii Per HKUCC 10837 NS DQ408559 NS
Stagonospora duoseptata Mas CBS 135093
HT
NS KF251758 NS
“S.”macropycnidia Len CBS 114202 GU296198 GU301873 GU349026
S. paludosa Mas CBS 135088
NT
NS KF251760 NS
S. (Neottiosporina)paspali Mas CBS 331.37 EU754073 EU754172 GU349079
S. perfecta Mas CBS 135099
HT
NS KF251761 NS
S. pseudocaricis Mas CBS 135132
HT
NS KF251763 NS
S. pseudopaludosa Mas CBS 136424
HT
NS KF777239 NS
S. pseudovitensis Mas S602
PT
NS KF251765 NS
S. trichophoricola Mas CBS 136764
HT
NS KJ869168 NS
S. uniseptata Mas CBS 135090
HT
NS KF251767 NS
Suttonomyces clematidis Mas MFLUCC 14-0240
HT
KP842920 KP842917 NS
Trematosphaeria grisea Tre CBS 332.50
HT
KF015641 KF015614 KF015698
T. grisea Tre CBS 135984 KF015632 KF015618 KF015694
T. pertusa Tre CBS 122368
ET
FJ201991 FJ201990 KF015701
1
Abbreviation of family names: Acr (Acrocalymmataceae), Bam (Bambusicolaceae), Dic (Dictyosporaceae), Did (Didymosphaeriaceae), IS (insertae sedis), Lat (Lator-
uaceae), Len (Lentitheciaceae), Lep (Leptosphaeriaceae), Lop (Lophiostomataceae), Mac (Macrodiplodiopsidaceae), Mas (Massarinaceae), Mor (Morosphaeriaceae), OG
(outgroup), Par (Parabambusicolaceae), Per (Periconiaceae), Ple (Pleosporaceae), Sul (Sulcatisporaceae), and Tre (Trematosphaeriaceae).
2
Isolate ex ET (epitype), HT (holotype), IT (isotype), LT (lectotype), NT (neotype), and PT (paratype).
3
NS: No sequence available in GenBank.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 83
Stagonospora perfecta KT 1726A
“Didymosphaeria” spartii CBS 183.58
Stagonospora tainanensis KT 1866
Stagonospora pseudovitensis S602 PT
Stagonospora perfecta CBS 135099 HT
Stagonospora pseudopaludosa CBS 136424 HT
Stagonospora uniseptata CBS 135090 HT
Stagonospora duoseptata CBS 135093 HT
Stagonospora (Neottiosporina) paspali CBS 331.37
Massarinaceae
Periconiaceae
Didymosphaeriaceae
Helminthosporium sp. yone 38
“Corynespopra” olivacea CBS 114450
Helminthosporium sp. H 4743
Helminthosporium sp. yone 63
Helminthosporium velutinum yone 96
Helminthosporium velutinum H 4626
Helminthosporium velutinum H 4739
Helminthosporium dalbergiae H 4628
Helminthosporium magnisporum H 4627 HT
“Corynespora” leucadendri CBS 135133 HT
Helminthosporium massarinum KT 1564 HT ★
Helminthosporium massarinum KT 838 PT ★
Byssothecium circinans CBS 675.92
Massarina eburnea CBS 473.64
Massarina eburnea H 3953
Massarina cisti CBS 266.62 HT
Stagonospora sp. KT 903
Stagonospora paludosa CBS 135088 NT
Stagonospora pseudocaricis CBS 135132 HT
Stagonospora trichophoricola CBS 136764 HT
Stagonospora pseudoperfecta KT 889 HT ★
Periconia pseudobyssoides H 4151
Periconia pseudobyssoides H 4790
Periconia byssoides H 4432
Periconia byssoides H 4600
Periconia byssoides H 4853
Periconia pseudodigitata KT 1395 HT ★
Periconia pseudodigitata KT 644 PT ★
Periconia pseudodigitata KT 1195A PT ★
Periconia digitata CBS 510.77
Periconia sp. KT 1820A
Periconia sp. KT 1825
Noosia banksiae CBS 129526 HT
“Sporidesmium” tengii HKUCC 10837
Periconia homothallica KT 916 HT ★
Periconia igniaria CBS 379.86
Periconia igniaria CBS 845.96
Spegazzinia deightonii yone 212
Spegazzinia deightonii yone 66
Spegazzinia sp. yone 279
Spegazzinia tessarthra SH 287
“Kalmusia (Montagnula)” anthostomoides CBS 615.86
Montagnula aloes CBS 132531 HT
Montagnula opulenta CBS 168.34
Munkovalsaria appendiculata CBS 109027 HT
Bimuria novae-zelandiae CBS 107.79 HT
Neokalmusia scabrispora KT 1023
Neokalmusia scabrispora KT 2202
Neokalmusia brevispora KT 1466
Neokalmusia brevispora KT 2313 ET
Deniquelata barringtoniae MFLUCC 110422 HT
Phaeodothis winteri CBS 182.58
“Paraconiothyrium” tiliae CBS 265.94 LT
Sporidesmiella fusiformis HKUCC 10831
Didymocrea sadasivanii CBS 438.65 IT
Alloconiothyrium aptrootii CBS 980.95 HT
Pseudocamarosporium brabeji CBS 119219 HT
Paracamarosporium leucadendri CBS 123027 HT
Pseudocamarosporium tilicola MFLUCC 13-0550 HT
Paracamarosporium psoraleae CBS 136628 HT
Pseudocamarosporium propinquum MFLUCC 13-0544 ET
Cucurbidothis pityophila CBS 149.32
Karstenula rhodostoma CBS 690.94
Karstenula rhodostoma CBS 691.94
Paraphaeosphaeria michotii CBS 652.86
Paraphaeosphaeria michotii KT 2222
Didymosphaeria rubi-ulmifolii MFLUCC 14-0023 HT
“Paraconiothyrium” fuckelii CBS 797.95
Paraconiothyrium estuarinum CBS 109850 HT
Paraconiothyrium maculicutis CBS 101461 HT
Letendraea eurotioides CBS 212.31
Letendraea helminthicola CBS 884.85
Letendraea padouk CBS 485.70
Kalmusia ebuli CBS 123120 NT
Kalmusia variispora CBS 121517 HT
Helminthosporium
sp
.
yone
38
“
Corynespopra
”
olivacea
CBS 11445
Suttonomyces clematidis MFLUCC 14-0240 HT
Periconia macrospinosa CBS 135663
Bambusistroma didymosporum MFLUCC 13-0862 HT
Flavomyces fulophazii CBS 135761 HT
Montagnula graminicola MFLUCC 13–0352 HT
“Paraphaeosphaeria” spartii MFLUCC 13–0398 HT
Pseudocamarosporium cotinae MFLUCC 14–0624 HT
Kalmusia spartii MFLUCC 14–0560 HT
Kalmusia italica MFLUCC 13–0066 HT
Paraconiothyrium thysanolaenae MFLUCC 10–0550 HT
88
54
82
89
73
95
90
58
73
96
81
100
52
97
54
99
66 67
80
76
100
98
51 99
99
100
90
95
55
97
99
100
100
68
98
99
83
73
100
81
63
89
62
51
68
99
69
99
57
58
97
98
73
87
77
100
96
85
95
78
51
Fig. 1. Maximum-likelihood tree of Massarineae based on the SSU and LSU nrDNA and tef1 regions. ML bootstrap proportion (BP) greater than 50 % is presented at the
nodes. An original isolate designation (or culture collection number) is noted after the species name. Sequences derived from holotype, isotype, neotype, paratype and epitype
TANAKA ET AL.
84
materials are indicated as HT, IT, NT, PTand ET, respectively. Species used for morphological observation in this study are formatted in bold. New names are marked by five-
pointed stars. Families, where known, are indicated with coloured blocks. The tree was rooted to Hysterobrevium mori and Hysterium pulicare in the Hysteriales. The scale bar
represents the number of nucleotide substitutions per site.
Gregarithecium curvisporum KT 922 HT ★
“Diplococcium” asperum CBS 139.95
Pseudocoleophoma polygonicola KT 731 HT ★
“Paraconiothyrium” flavescens CBS 178.93
Dendryphiella vinosa
Dictyosporium inflatum NTOU 3855
Dictyosporium toruloides CBS 209.65
Digitodesmium bambusicola CBS 110279 HT
Dictyosporium pseudomusae yone 234 HT ★
Dictyosporium pseudomusae KH 412 PT ★
Dictyosporium aff. heptasporum KH 332
Pseudodictyosporium elegans CBS 688.93 HT
Pseudodictyosporium wauense NBRC 30078
Dictyosporium bulbosum yone 221
Dictyosporium strelitziae CBS 123359 HT
Dictyosporium hughesii KT 1847
Dictyosporium aff. bulbosum KH 375
Dictyosporium tetrasporum KT 2865
Dictyosporium stellatum CCFC 241241 HT
Dictyosporium digitatum KH 401
Dictyosporium digitatum yone 280
Dictyosporium digitatum KT2660
Dictyosporium elegans NBRC 32502
Dictyosporium alatum ATCC 34953 HT
Aquaticheirospora lignicola RK-2006a HT
Katumotoa bambusicola KT 1517a PT
Neoophiosphaerella sasicola KT 1706 ET ★
Tingoldiago graminicola KH 68 HT
Tingoldiago graminicola KT 891 PT
Tingoldiago graminicola KH 155 PT
Setoseptoria magniarundinacea KT 1174 HT ★
“Stagonospora” macropycnidia CBS 114202
Setoseptoria arundinacea CBS 619.86
Setoseptoria arundinacea CBS 123131
Setoseptoria arundinacea KT 552 ★
Setoseptoria arundinacea KT 600 ★
Lentithecium aquaticum CBS 123099 HT
Lentithecium clioninum KT 1149A HT ★
Lentithecium clioninum KT 1220 PT ★
Lentithecium fluviatile CBS 122367
Lentithecium pseudoclioninum KT 1111 PT ★
Lentithecium pseudoclioninum KT 1113 HT ★
Keissleriella breviasca KT 540 PT ★
Keissleriella breviasca KT 581 PT ★
Keissleriella breviasca KT 649 HT ★
Keissleriella culmifida KT 2642
Keissleriella culmifida KT 2308
Keissleriella sp. KT 895
Keissleriella quadriseptata KT 2292 HT ★
Keissleriella gloeospora KT 829
Keissleriella taminensis KT 571
Keissleriella taminensis KT 678
Keissleriella taminensis KT 594
Keissleriella cladophila CBS 104.55
Keissleriella genistae CBS 113798
Keissleriella linearis IFRD 2008
Keissleriella yonaguniensis KT 2604 HT ★
Pseudocoleophoma calamagrostidis KT 3284 HT ★
Setoseptoria phragmitis CBS 114966 PT
Setoseptoria phragmitis CBS 114802 HT
Keissleriella trichophoricola CBS 136770 HT
Keissleriella poagena CBS 136767 HT
Pleurophoma pleurospora CBS 130329 LT
Dictyosporiaceae
Lentitheciaceae
Unknown Clade I
Fuscostagonospora sasae KT 1467 HT ★
Dictyosporium thailandicum MFLUCC 13-0773 HT
Dictyosporium meiosporum MFLUCC 10-0131 HT
Keissleriella sparticola MFLUCC 14–0196 HT
Phragmocamarosporium platani MFLUCC 14-1191 HT
Phragmocamarosporium hederae MFLUCC 13-0552 HT
Murilentithecium clematidis MFLUCC 14-0561 HT
Darksidea alpha CBS 135650 HT
Darksidea delta CBS 135638 HT
Darksidea gamma CBS 135634 HT
Darksidea beta CBS 135637 HT
Darksidea zeta CBS 135640 HT
Darksidea epsilon CBS 135658 HT
Poaceascoma helicoides MFLUCC 11-0136 HT
82
74
52
97
90
54
77
100
75
83
94 87
51
100 100
76
63 100
63
96
99
7678
96
54
69 65
96
80
99
91
98
100
59
64
83
95
84
90
83
67
82
89
79
85
100
74
71
62
68
71
62
Fig. 1. (Continued).
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 85
freshwater environments. The genus is characterised by pig-
mented, cheiroid conidia formed on sporodochial colonies. It has
been considered that the genus is closely related to the Mas-
sarinaceae in the Pleosporales based on phylogenetic analysis
using SSU and LSU nrDNA sequences (Tsui et al. 2006). After
taxonomic revision of the genus (Goh et al. 1999, Cai et al.
2003), 33 species were accepted as Dictyosporium (Crous
et al. 2009a). In addition, 16 species have been further recor-
ded in this genus (Manoharachary et al. 2007, Zhang et al.
2009a, Wongsawas et al. 2009, Hu et al. 2010, McKenzie
2010, Crous et al. 2011a, Whitton et al. 2012, Kirschner et al.
2013, Liu et al. 2015, Prasher & Verma 2015).
Dictyosporium bulbosum Tzean & J.L. Chen, Mycol. Res. 92:
500. 1989. Fig. 2A–C.
Specimen examined:Japan, Okinawa, Isl. Iriomote, near Maryudu-falls, on dead
twigs of woody plant, 27 Sep. 2007, H. Yonezawa & K. Tanaka, yone 221 = HHUF
29990, culture MAFF 243835.
Leptosphaeria doliolum CBS 505.75
Pleospora herbarum CBS 191.86
Polyschema larviformis CBS 463.88
Polyschema congolensis CBS 542.73 HT
Polyschema terricola CBS 301.65 HT
Magnicamarosporium iriomotense KT 2822 HT ★
Sulcatispora berchemiae KT 1607 HT ★
Bambusicola bambusae MFLUCC 11-0614 HT
Bambusicola massarinia MFLUCC 11-0389 HT
Bambusicola irregulispora MFLUCC 11-0437 HT
Bambusicola splendida MFLUCC 11-0439 HT
Monodictys sp. JO 10
Monodictys sp. KH 331
Aquastroma magniostiolata KT 2485 HT ★
Parabambusicola bambusina H 4321 ★
Parabambusicola bambusina KH 139 ★
Parabambusicola bambusina KT 2637 ★
Macrodiplodiopsis desmazieri CBS 221.37
Halomassarina thalassiae BCC 17054
Halomassarina thalassiae JK 5262D
Trematosphaeria pertusa KT 1496
Trematosphaeria pertusa CBS 122368 ET
Falciformispora lignatilis BCC 21117
Bactrodesmium cubense CBS 680.96
Aquilomyces rebunensis KT 732 HT ★
Clypeoloculus towadaensis KT 1340 HT ★
Clypeoloculus microsporus KT 1131 HT ★
Clypeoloculus akitaensis KT 788 HT ★
Clypeoloculus hirosakiensis KT 1283 HT ★
Helicascus thalassioideus KH 242
Helicascus thalassioideus CBS 110441
Helicascus thalassioideus MFLUCC10-0911
Helicascus nypae BCC 36751
Helicascus nypae BCC 36752
Morosphaeria ramunculicola KH 220
Morosphaeria ramunculicola BCC 18404
Morosphaeria velatispora BCC 17059
Morosphaeria velatispora KH 221
Morosphaeria velatispora KH 218
Inflatispora pseudostromatica CBS 123110 HT
Monodictys capensis HR 1
Asteromassaria pulchra CBS 124082
Trematosphaeria pertusa KT 3315
Trematosphaeria pertusa KT 3314
Morosphaeriaceae
Sulcatispora acerina KT 2982 HT ★
Camarographium koreanum CBS 117159 HT
Pseudochaetosphaeronema larense CBS 640.73 HT
Trematosphaeria grisea CBS 332.50 HT
Trematosphaeria grisea CBS 135984
Falciformispora senegalensis CBS 196.79 HT
Falciformispora tompkinsii CBS 200.79 HT
“Pithomyces” valparadisiacus CBS 113339
Helicascus aegyptiacus FWCC 99 HT
Helicascus elaterascus HKUCC 7769
Helicascus aquaticus MFLUCC 10-0918 HT
Helicascus aquaticus KT1544
Helicascus elaterascus KT2673
Helicascus elaterascus KT2682
Lophiostoma arundinis JCM 13550
Lophiostoma macrostomum JCM 13544
Hysterobrevium mori CBS 12356
Hysterium pulicare CBS 12337
Trematosphaeriaceae
Bambusicolaceae
Parabambusicolaceae
Unknown Clade II
Unknown Clade III
Unknown Clade IV
Latoruaceae
Macrodiplodiopsidaceae
Pseudoxylomyces elegans KT 2887
0.03
Latorua caligans CBS 576.65 HT
Neobambusicola strelitziae CBS 138869 HT
Latorua grootfonteinensis CBS 369.72 HT
Bambusicola loculata MFLUCC 13-0856 HT
Palmiascoma gregariascomum MFLUCC 11–0175 HT
Multiseptospora thailandica MFLUCC 11–0183 HT
Macrodiplodiopsis desmazieri CBS 140062 ET
Aquilomyces patris CBS 135661 HT
Helicascus unilocularis MJF14020 HT
PLEOSPORALES
x2
MASSARINEAE
Unknown Clade V
89
100
81
64
71
87
94
58
97
97 74
99
99
100
92
72
85
98
79
100
97
99
96
99
94
95
93
100
79
100
73
88
96 98
72
79
99
100 98
100
97
79
100
83 73
52
63
61
98
100
99
65
92
100
91
100
100
Sulcatisporaceae
63
Fig. 1. (Continued).
TANAKA ET AL.
86
Note: This specimen was identified as D. bulbosum and
morphological features were described and illustrated by
Hirayama et al. (2012).
Dictyosporium aff. bulbosum Fig. 2D–F.
Specimen examined:Japan, Okinawa, Isl. Iriomote, Uehara, near Tropical
Biosphere Research Center (Ryukyu Univ.), small stream, on submerged twigs of
woody plant, 13 Jul. 2011, K. Hirayama & K. Tanaka, KH 375 = HHUF 30127,
culture JCM 19403 = MAFF 243829.
Notes: The above isolate produced an asexual morph in culture,
which is similar to that on the natural specimen. Conidia in
culture were 36–50 × 26–33 μm (av. 45.7 × 29.1 μm, n = 33), l/w
1.3–1.8 (av. 1.6, n = 33), consisting of 29–47 cells arranged in
5–6 rows, with apical appendages. These features almost agree
with the description in the protologue of D. bulbosum (Tzean &
Chen 1989), but the ITS sequences of our material (KH 375;
GenBank LC014542) differed from those of D. bulbosum (Gen-
Bank LC014544 and DQ018086) in ca. 8 % (40/515) positions.
Dictyosporium digitatum J.L. Chen et al., Mycol. Res. 95:
1145. 1991. Fig. 2G–I.
Specimens examined:Japan, Okinawa, Isl. Ishigaki, trail of Mt. Omoto, on dead
stems of herbaceous plant, 16 Jul. 2011, K. Hirayama & K. Tanaka, KH
401 = HHUF 30128, culture JCM 19404 = MAFF 243830; Okinawa, Isl. Iriomote,
Komi, on dead wood of Castanopsis sieboldii, 15 Sep. 2009, Y. Kurihara, KT
2660 = HHUF 30131, culture JCM 19405 = MAFF 243833; Okinawa, Isl. Iriomote,
Kanpire-falls, on dead twigs of woody plant, 21 Nov. 2008, K. Tanaka & K. Hir-
ayama, yone 280 = HHUF 30093, culture MAFF 243837.
Notes: The morphological characters of our specimens are
consistent with those of D. digitatum (Chen et al. 1991), as re-
ported by Hirayama et al. (2012). The ITS sequences of this
species from our three isolates were highly similar
(99.1–99.8 %) but their similarities with a deposited sequence of
D. digitatum in GenBank (GenBank DQ018089) were rather low
(90.1–90.4 %).
Dictyosporium aff. heptasporum Fig. 2J–L.
Specimen examined:Japan, Okinawa, Isl. Ishigaki, Mt. Banna, near small
stream, on dead twigs of woody plant, 14 Jul. 2011, K. Hirayama & K. Tanaka, KH
332 = HHUF 30126, culture JCM 19406 = MAFF 243828.
Notes: The morphological features of the above specimen were
as follows; conidia 70–90 × 21–31 μm (av. 79.7 × 23.3 μm,
AD
LKI
F
E
CB
JG
NOS
TUV
MPQRW
XY
H
Fig. 2. Dictyosporium spp. A–C. D. bulbosum (A from culture yone 221; B, C from yone 221); D –F. D. aff. bulbosum (culture KH 375); G–I. D. digitatum (G from culture KT
2660; H from culture yone 280; I from culture KH 401); J–L. D. aff. heptasporum (KH 332); M–O. D. hughesii (M, O from KT 1847; N from culture KT 1847); P –V.
D. pseudomusae (P from yone 234; Q, T from culture yone 234; R, U from culture KH 412; S, V from KH 412); W–Y. D. tetrasporum (culture KT 2865). A, D, G, J, M, P –R, W.
Conidial masses on culture media or natural substrates (A, D, G, Q, R, W from culture; J, M, P from nature); B, C, E, F, H, I, K, L, N, O, S–V, X, Y. Conidia (arrowheads indicate
conidial appendages). Scale bars: A, D, G, J, M, P–R, W = 200 μm; B, C, E, F, H, I, K, L, N, O, S–V, X, Y = 10 μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 87
n = 43), cylindrical, l/w 3.1–3.9 (av. 3.4, n = 43), with (5–)7 rows,
16–18-septate, without appendages. This agrees with the details
in the description of D. heptasporum (Goh et al. 1999), but the
conidia in our material are somewhat larger (vs.
50–80 × 20–30 μm; Goh et al. 1999). A BLAST search using ITS
sequences from our culture showed D. heptasporum (GenBank
DQ018090) as the closest species, but the similarity was rela-
tively low (493/518 = 95.2 %).
Dictyosporium hughesii McKenzie, Mycotaxon 111: 156. 2010.
Fig. 2M–O.
Specimen examined:Japan, Kagoshima, Isl. Yakushima, Shirataniunsuikyo, on
dead twigs of Stewartia monadelpha, 18 Oct. 2005, K. Tanaka & T. Hosoya, KT
1847 = HHUF 30130 = TNS-F-12407, culture JCM 19407 = MAFF 243832.
Note: This collection was identified as D. hughesii (McKenzie
2010) based on the conidial morphology; conidia
43–51 × 18–28 μm (av. 46.8 × 24.8 μm, n = 30), l/w
1.6–2.3(–2.6) (av. 1.9, n = 30), consisting of 50–71 cells ar-
ranged in 6–7 rows, with or without apical appendages.
Dictyosporium pseudomusae Kaz. Tanaka, G. Sato & K.
Hiray., sp. nov. MycoBank MB811297. Fig. 2P–V.
Etymology: After its morphological similarity to Dictyosporium
musae.
Sporodochia on natural substrate scattered, punctiform, dark
brown to black. Mycelium immersed, 170–490 μm diam. Co-
nidiophores micronematous, not differentiated from vegetative
hyphae. Conidiogenous cells holoblastic, cylindrical, 5.5–8μm
wide. Conidia solitary, brown, ellipsoid to cylindrical, cheiroid, not
complanate, (58–)61–78(–81) × 19–29(–33) μm (av.
69.9 × 22.9 μm, n = 60), l/w (2.2–)2.5–3.8(–4.0) (av. 3.1,
n = 60), consisting of 78–100 cells arranged in (6–)7 rows and
basal connecting cell (6–8 × 5.5–8μm); each row cylindrical,
with 13–15 cells. Appendages globose to subglobose, hyaline,
6–11.5 μm diam, bearing from apical cells or side of outer rows.
Sexual morph unknown.
Colonies on PDA (after 4 wk) attaining a diam of 3.9–4.4 cm,
white to rosy buff; reverse buff to cinnamon; rosy vinaceous
pigment produced. In culture asexual morph formed.
Specimens examined:Japan, Okinawa, Isl. Ishigaki, Mt. Banna, near small
stream, on dead twigs of Bamboo, 14 Jul. 2011, K. Hirayama & K. Tanaka, KH
412 = HHUF 30129, culture JCM 19408 = MAFF 243831; Okinawa, Isl. Iriomote,
Inamori-path, on dead twigs of woody plant, 25 Sep. 2007, H. Yonezawa & K.
Tanaka (holotype yone 234 = HHUF 30133, culture ex-type CBS 139686 = JCM
19409 = MAFF 243836).
Notes: Among the described species, D. pseudomusae is most
similar to D. musae (Photita et al. 2002) in having large-sized
conidia comprised of seven cell rows and with appendages
arising from side cells of the outer rows. Conidia of the latter
species, however, are smaller (45–65 × 20 –27 μm, av.
55.9 × 23.5 μm) and comprised of fewer cells (49–77 cells;
Photita et al. 2002). The closest species to D. pseudomusae
appear to be D. digitatum [GenBank DQ018089; identities = 504/
516 (97.7 %), gaps = 2/516] and D. giganticum [GenBank
DQ018095; identities = 497/516 (96.3 %), gaps = 2/516] from a
BLAST search using ITS sequences.
Dictyosporium tetrasporum L. Cai & K.D. Hyde, Mycoscience
48: 290. 2007. Fig. 2W–Y.
Specimen examined:Japan, Okinawa, Isl. Ishigaki, Mt. Banna, near small
stream, on dead twigs of woody plant, 16 Jul. 2011, K. Tanaka & K. Hirayama, KT
2865 = HHUF 30132, culture JCM 19410 = MAFF 243834.
Notes: The collection matches the original description of
D. tetrasporum (Cai & Hyde 2007a). The characters of our
specimen are as follows; conidia 22–37 × 15–21 μm (av.
27.5 × 17.5 μm, n = 30), l/w 1.2–2.2 (av. 1.6, n = 30), consisting
of 11–28 cells arranged in (3–)4 rows, without apical append-
ages. In culture, conidia were produced that were identical to
those on the natural specimen (24–38 × 16–24 μm; av.
30.3 × 20.1 μm, n = 32).
Gregarithecium Kaz. Tanaka & K. Hiray., gen. nov. MycoBank
MB811298.
Etymology: Referring to the gregarious ascomata.
Ascomata grouped, immersed to erumpent, depressed globose
to hemispherical with flattened base in section. Ostiolar neck
short terete, central, with periphyses, covered by black clypeus.
Ascomatal wall composed of thin-walled cells, surrounded by
vertically-orientated stromatic tissue. Pseudoparaphyses
septate, branched and anastomosed. Asci fissitunicate, cylin-
drical, short-stalked, with 8 biseriate ascospores. Ascospores
broadly fusiform, with a median septum, hyaline, smooth, sur-
rounded by an entire sheath. Asexual morph unknown.
Type species:Gregarithecium curvisporum Kaz. Tanaka &
K. Hiray.
Notes: In its depressed globose ascomata with clypeate ostiole,
Gregarithecium is reminiscent of species in Massarina s. lat.
(Aptroot 1998, Tanaka & Harada 2003b), but Gregarithecium can
be separated from the latter by the presence of stromatic tissue
surrounding the ascomata. Massarina s. lat. is regarded as being
polyphyletic and thus has recently been segregated into several
new genera. These are scattered in the Lentitheciaceae (Tin-
goldiago;Hirayama et al. 2010), Morosphaeriaceae (Moro-
sphaeria;Suetrong et al. 2009), Trematosphaeriaceae
(Halomassarina;Suetrong et al. 2009), Tetraplosphaeriaceae
(Triplosphaeria;Tanaka et al. 2009), and Lindgomycetaceae
(Lindgomyces;Hirayama et al. 2010), but are not so far reported
in the Dictyosporiaceae.
Gregarithecium curvisporum Kaz. Tanaka & K. Hiray.,
sp. nov. MycoBank MB811299. Fig. 3.
Etymology: From the Latin curvi-, in reference to the curved
ascospores.
Ascomata grouped in numbers of 3–6, immersed to erumpent,
depressed globose to hemispherical with flattened base in sec-
tion, 140–180 μm high, 290–430 μm diam. Ostiolar neck short
terete, central, with an ostiole (25–50 μm diam) and short
periphyses, covered by a black clypeus composed of small cells
(2.5–7.5 × 1–3μm). Ascomatal wall in longitudinal section
8–12 μm thick at the sides, composed of thin-walled, flattened,
pale brown cells, surrounded by vertically-oriented stromatic
TANAKA ET AL.
88
tissue composed of rectangular to polygonal cells
(5–25 × 5–16 μm) or by compact brown hyphae. Pseudopar-
aphyses septate, branched and anastomosed, 2–3.5(–5) μm
wide. Asci fissitunicate, cylindrical, 73.5–102 × 11–15.5 μm (av.
87.4 × 13.6 μm, n = 50), rounded at the apex, with a shallow
ocular chamber (0.5–1μm high), short-stalked (5–15 μm long;
av. 9.6 μm, n = 32), with 8 biseriate ascospores. Ascospores
broadly fusiform, mostly curved, 19–31 × 4.5–6.5 μm (av.
24.2 × 5.8 μm, n = 50), l/w (3.3–)3.6–4.9(–5.2) (av. 4.2, n = 50),
with a median (0.50) septum and constricted, hyaline, smooth,
with an entire sheath; sheath sharply delimited and 1–2μm wide
at first, diffuse and 4–8μm wide at a later. Senescent asco-
spores 3-septate, thick-walled, dark brown.
Colonies on PDA (after 4 wk) attaining a diam of 6.2–6.8 cm, buff
to smoke grey; reverse pale luteous to sienna; no pigment
produced. In culture ascomatal and spermatial morphs observed.
Asci 82.5–101.5 × 14.5–17.5 μm. Ascospores 21–31 × 6–8μm
(av. 24.5 × 7.0 μm, n = 31), l/w 3.1–4.2 (av. 3.5, n = 31).
Spermatia 2.2–3.5 × 1.4–2.1 μm (av. 3.0 × 1.8 μm, n = 20),
subglobose, hyaline.
Specimen examined:Japan, Aomori, Hirosaki, Kozawa, on dead culms of Sasa
sp., 25 Oct. 2002, T. Handa (holotype KT 922 = HHUF 30134, culture ex-type
CBS 139688 = JCM 19411 = MAFF 243838).
Pseudocoleophoma Kaz. Tanaka & K. Hiray., gen. nov.
MycoBank MB811300.
Etymology: Referring to the similarity of the asexual morph with
that of Coleophoma.
Ascomata scattered to grouped, immersed to erumpent,
globose to subglobose in section. Ostiolar neck central,
composed of subglobose dark brown cells. Ascomatal wall
composed of polygonal to rectangular cells. Pseudoparaphyses
numerous. Asci fissitunicate, cylindrical to clavate, short-
stalked, with 8 ascospores. Ascospores fusiform, 1-septate,
smooth, with a conspicuous sheath. Conidiomata coleophoma-
like, pycnidial, subglobose. Conidiophores absent. Con-
idiogenous cells phialidic, doliiform to lageniform. Conidia cy-
lindrical, hyaline, smooth.
Type species:Pseudocoleophoma calamagrostidis Kaz. Tanaka
& K. Hiray.
Notes: The pycnidial asexual morph of Pseudocoleophoma
bears a slight resemblance to that of Coleophoma. The type
species of Coleophoma (C. crateriformis), however, has pycnidia
possessing paraphyses that are not found in Pseudocoleo-
phoma, and is a member of the Dothideales, rather than the
Pleosporales (De Gruyter et al. 2009).
Pseudocoleophoma calamagrostidis Kaz. Tanaka & K. Hiray.,
sp. nov. MB811301. Fig. 4.
Etymology: Referring to the host plant.
Ascomata scattered, immersed, erumpent at the top, globose to
depressed globose, 140–200 μm high, 160–220 μm diam,
ostiolate. Ostiolar neck central. Ascomatal wall in longitudinal
section 5–10 μm thick at sides, composed of 2–3 layers of thin-
walled, polygonal, flattened, 5–12 × 2–5μm, brown cells; of
polygonal to subglobose, 2–5 × 1.5–2.5 μm cells around ostiole.
Pseudoparaphyses septate, branched and anastomosed,
2.5–5μm wide. Asci fissitunicate, cylindrical,
62.5–80 × 7.5–10 μm (av. 69.0 × 8.4 μm, n = 50), rounded at the
apex, with a shallow ocular chamber, short-stalked (5–12 μm
long), with 8 biseriate ascospores. Ascospores narrowly fusiform,
L
J
G
FE
D
AB C
K
H
I
Fig. 3. Gregarithecium curvisporum. A, B. Ascomata on the natural host surface; C. Ascomata in culture (on rice straw); D, E. Ascomata in longitudinal section; F. Ascomatal
wall; G. Pseudoparaphyses; H. Ascus; I–K. Ascospores (arrowheads indicate mucilaginous sheath); L. Spermatia. A, B, D–I from KT 922; C, J –L from culture KT 922. Scale
bars: A = 2 mm; B, C = 500 μm; D, E = 100 μm; F–L=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 89
(14.5–)16–19(–21) × 3–4.5 μm (av. 17.4 × 3.8 μm, n = 30), l/w
4.0–5.3 (av. 4.5, n = 30), with a nearly median (0.43–0.54, av.
0.50, n = 26) septum, constricted at the septum, hyaline, smooth,
with an entire sheath; sheath sharply delimited, 1–2μm wide at
sides, 4–6μm long at both ends, staining with Black-Blue ink
when in fresh condition.
Colonies on PDA (after 4 wk) attaining a diam of 2.3–3.2 cm, buff
to honey; reverse similar; no pigment produced. In culture coe-
lomycetous asexual morph formed. Conidiomata pycnidial,
220–300 μm high, (150–)250–500 μm diam, immersed to
erumpent, depressed globose. Ostiolar neck well-developed,
75–100 μm long, 85–100 μm wide, cylindrical, central. Con-
idiomatal wall in longitudinal section uniformly 7.5–15 μm thick,
composed of 3–4 layers of slightly thick-walled, polygonal to
subglobose, 6–15 × 2–6.5 μm, pale brown cells; of polygonal to
rectangular, 2–6 × 1.5 –2.5 μm cells around ostiole. Co-
nidiophores absent. Conidiogenous cells phialidic,
5–9×2–4μm, doliiform to subglobose. Conidia cylindrical,
aseptate, hyaline, smooth, 6–10 × 2–2.5 μm (av. 8.6 × 2.2 μm,
n = 50), l/w 2.9–4.7 (av. 3.9, n = 50).
Specimen examined:Japan, Aomori, Hirosaki, Mt Iwaki, on dead leaves of
Calamagrostis matsumurae, 27 Jul. 2013, K. Tanaka (holotype KT 3284 = HHUF
30450, culture ex-type CBS 139700).
Note: This species is phylogenetically close to P. polygonicola,
but the ITS sequence similarity between these two species is
relatively low (490/521 = 94.0 %).
Pseudocoleophoma polygonicola Kaz. Tanaka & K. Hiray.,
sp. nov. MycoBank MB811302. Fig. 5.
Etymology: Referring to the host plant.
Ascomata scattered to 2–4 grouped, immersed to erumpent,
230–310 μm high, 280–350 μm diam. Ostiolar neck central,
50–75 μm long, 70–100 μm wide. Ascomatal wall in longitudinal
section 7–13 μm thick at sides. Pseudoparaphyses 2–2.5 μm
wide. Asci cylindrical to clavate, (67–)74–90(–100) ×
9–12.5 μm (av. 81.3 × 10.7 μm, n = 48). Ascospores fusiform,
(17.5–)19–23(–25) × 4–6μm (av. 20.6 × 4.8 μm, n = 50), l/w
3.6–4.8 (av. 4.2, n = 45), with a septum supramedian
(0.46–0.50; av. 0.48, n = 40), surrounded by a sheath; sheath
1–2μm wide at sides, 2–5μm long at both ends, staining with
Black-Blue ink when in fresh condition.
Colonies on PDA (after 4 wk) attaining a diam of 3.4–4.8 cm,
white; reverse white to straw; no pigment produced. In culture
coelomycetous asexual morph formed. Conidiomata pycnidial,
AB
CD
E
FGH
IJKL
RS
Q
M
NO P
Fig. 4. Pseudocoleophoma calamagrostidis. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall at side; E. Ascomatal wall
surface around ostiole; F. Pseudoparaphyses; G, H. Asci; I–K. Ascospores (arrowheads indicate gelatinous sheath; I. in Black-Blue ink); L. Germinating ascospore; M, N.
Conidiomata in culture (on rice straw); O. Conidioma in longitudinal section; P. Conidiomatal wall at side; Q. Conidiomatal wall around ostiole; R. Conidiogenous cell; S. Conidia.
A–L from KT 3284; M–S from culture KT 3284. Scale bars: A, B, M, N = 500 μm; C, E, O = 50 μm; D, F–L, P–S=10μm.
TANAKA ET AL.
90
170–250 μm diam. Conidiomatal wall in longitudinal section
12–15 μm wide at sides. Conidiophores absent. Conidiogenous
cells phialidic, 7–17 × 3.5–5μm, doliiform to lageniform, formed
all around the locular cavity. Conidia cylindrical, aseptate, hya-
line, smooth, (9–)11.5–18(–21.5) × 3–4.5 μm (av.
14.4 × 3.4 μm, n = 31).
Specimen examined:Japan, Hokkaido, Isl. Rebun, Kafuka, Nairo-river, on dead
stems of polygonaceous plant, 30 Aug. 2001, K. Tanaka (holotype KT
731 = HHUF 27558, culture ex-type CBS 139701 = JCM 19412 = MAFF
239468).
Notes: Additional details of this species were reported by Tanaka
& Harada (2003b), who misidentified this fungus as Massarina
rubi based mostly on the morphology of the sexual morph.
However, Pseudocoleophoma is phylogenetically different from
M. rubi, which belongs to the Lophiotremataceae (Zhang et al.
2009b, Hirayama & Tanaka 2011b). This species resembles
P. calamagrostidis but has larger ascospores (20.6 × 4.8 μmvs.
17.4 × 3.8 μm) and conidia (14.4 × 3.4 μmvs. 8.6 × 2.2 μm).
Didymosphaeriaceae Munk, Dansk Bot. Ark. 15 (2): 128. 1953.
=Montagnulaceae M.E. Barr, Mycotaxon 77: 194. 2001.
Type genus:Didymosphaeria Fuckel.
Karstenula Speg., Decades Mycologicae Italicae 7–12: no. 94.
1879.
Type species:Karstenula rhodostoma (Alb. & Schwein.) Speg.
Notes: This genus is phylogenetically close to Para-
phaeosphaeria, but can be distinguished from the latter by the
ascomata surrounded by well-developed subiculum and cylin-
drical asci with uniseriate ascospores.
Karstenula rhodostoma (Alb. & Schwein.) Speg., Decades
Mycologicae Italicae 7–12: no. 94. 1879. Fig. 6.
Basionym:Sphaeria rhodostoma Alb. & Schwein., Consp. Fung.
(Leipzig): 43. 1805.
FE
DC
BA
Fig. 6. Karstenula rhodostoma. A. Ascoma in longitudinal section; B. Ascomatal wall; C. Pseudoparaphyses; D. Ascus; E, F. Ascospores. All from UPS (F-141152) 425947.
Scale bars: A = 50 μm; B–F=10μm.
L
E
D
J
I
G
O
N
M
K
H
F
AB
C
Fig. 5. Pseudocoleophoma polygonicola. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus;
G–J. Ascospores (arrowheads indicate gelatinous sheath; J. in Black-Blue ink); K. Conidiomata in culture; L. Conidioma in longitudinal section; M. Conidiomatal wall; N.
Conidiogenous cells; O. Conidia. A–J from KT 731; K–O from culture KT 731. Scale bars: A = 2 mm; B, K = 500 μm; C, L = 50 μm; D –J, M –O=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 91
Asci cylindrical, 145–175 × 12.5–14.5 μm (av. 162.2 × 13.4 μm,
n = 20), with a short stipe (14–23 μm long). Ascospores cylin-
drical to ellipsoid, 22–26 × 8–10 μm (av. 23.7 × 9.1 μm, n = 20),
l/w 2.3–2.9 (av. 2.6, n = 20), with a submedian primary septum
(0.50–0.54; av. 0.52, n = 20), with 3 transverse septa (1+1+1),
with or without 1 longitudinal septum.
Specimen examined:Sweden,“Odlingen”field, ca. 250 m S of Jerusalem, on
twigs of Frangula alnus, Jan. 1995, K. Holm & L. Holm, UPS (F-141152) 425947,
culture CBS 691.94.
Notes: For further description of this species including its asexual
morph (Microdiplodia frangulae), see Constantinescu (1993) and
Zhang et al. (2012). More than 360 species are recorded as
Microdiplodia (MycoBank, http://www.mycobank.org), but the
validity of this genus is unknown due to lack of phylogenetic
information regarding the lectotype species, M. conigena
(Clements & Shear 1931).
Neokalmusia Kaz. Tanaka et al., Fungal Divers. 68: 92. 2014.
Type species:Neokalmusia brevispora (Nagas. & Y. Otani) Kaz.
Tanaka et al.
Notes:Neokalmusia is characterised by subglobose to oblong
ascomata including several pseudothecia in a row and verrucose
ascospores (Ariyawansa et al. 2014). Two bambusicolous
species, N. brevispora on Sasa and N. scabrispora on Phyllos-
tachys are known in this genus. Although they share many
morphological features, monophyly of Neokalmusia was not
supported in this study (Fig. 1). Additional taxa and sequence data
of this genus are needed to evaluate validity of Neokalmusia.
Neokalmusia brevispora (Nagas. & Y. Otani) Kaz. Tanaka
et al., Fungal Divers. 68: 92. 2014. Fig. 7.
Basionym:Phaeosphaeria arundinacea var. brevispora Nagas. &
Y. Otani, Rep. Tottori Mycol. Inst. 15: 38. 1977.
Specimens examined:Japan, Hokkaido, Sapporo, Moiwa-yama, on culms of Sasa
sp., 13 Jun. 1972, E. Nagasawa (holotype of Phaeosphaeria arundinacea var.
brevispora TMI 3175); Fukushima, Minamiaizu, Ose pond, on dead twigs of Sasa
sp., 30 Aug. 2003, N. Asama, KT 1466 = HHUF 28229, culture CBS 120248 = JCM
13543 = MAFF 239276; Hokkaido, Isl. Rishiri, Afutoromanai-river, on dead twigs of
Sasa kurilensis, 29 Jul. 2007, K. Tanaka & G. Sato (epitype designated here KT
2313 = HHUF 30016, MBT202863, culture ex-epitype NBRC 106240).
Notes: For other descriptions of this species, see Nagasawa &
Otani (1977), Shoemaker & Babcock (1989), Tanaka & Harada
(2004), and Ariyawansa et al. (2014). Although this species
was transferred to Kalmusia (Zhang et al. 2009b), subsequent
molecular studies (Hyde et al. 2013, Zhang et al. 2014a) did not
support this placement, and thus Neokalmusia was established
to accommodate N. brevispora as the type species of this genus
(Ariyawansa et al. 2014). We here designate an epitype spec-
imen for this species.
C
A
EF
G
J
H
D
B
I
Fig. 7. Neokalmusia brevispora. A. Ascomata on the natural host surface; B, C. Ascomata in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G–I.
Ascospores (arrowheads indicate gelatinous sheath; I. in India ink); J. Germinating ascospore. A, F, G, J from KT 2313; B–D, I from KT 1466; E, H from culture KT 1466. Scale
bars: A = 1 mm; B, C = 100 μm; D–J=10μm.
C
G
DE
B
A
I
F
H
J
Fig. 8. Neokalmusia scabrispora. A. Ascomata on the natural host surface; B, C. Ascomata in longitudinal section (B. in lactophenol cotton blue); D. Ascomatal wall; E.
Pseudoparaphyses; F. Ascus; G, H. Ascospores (arrowheads indicate gelatinous sheath; H. in India ink); I. Germinating ascospores; J. Spermatia. A–D from KT 1023; E–I from
KT 2202; J from culture KT 1023. Scale bars: A = 1 mm; B, C = 100 μm; D–J=10μm.
TANAKA ET AL.
92
Neokalmusia scabrispora (Teng) Kaz. Tanaka et al., Fungal
Divers. 68: 92. 2014. Fig. 8.
Basionym:Leptosphaeria scabrispora Teng, Sinensia, Shanghai
4: 378. 1934.
Specimens examined:Japan, Tochigi, Kanuma, near Ooashi-river, on dead twigs
of Phyllostachys bambusoides, 6 Mar. 2003, N. Asama, KT 1023 = HHUF 28608,
culture CBS 120246 = JCM 12851 = MAFF 239517; Kagoshima, Kumagegun, Isl.
Yakushima, Miyanoura-river, riverbank, on dead twigs of Phyllostachys bambu-
soides, 17 Mar. 2007, K. Tanaka & H. Yonezawa, KT 2202 = HHUF 30013,
culture NBRC 106237.
Notes: For a description of this species, see Tanaka et al.
(2005a). This fungus was originally described as a Lep-
tosphaeria (Teng 1934), and later transferred to Massariosphaeria
(Shoemaker & Babcock 1989)orKalmusia (Tanaka et al. 2005a).
Molecular studies, however, did not support these placements
(Tanaka et al. 2009, Zhang et al. 2014a). This species is currently
treated as Neokalmusia (Ariyawansa et al. 2014), but we were not
able to confirm the congenericity of N. scabrispora with
N. brevispora (type species of Neokalmusia)(Fig. 1).
Paraphaeosphaeria O.E. Erikss., Ark. Bot. 6: 405. 1967.
Type species:Paraphaeosphaeria michotii (Westend.) O.E. Erikss.
Notes: The genus Paraphaeosphaeria was established by
Eriksson (1967) as a generic segregate from Leptosphaeria.To
date, about 35 taxa have been described as species within
Paraphaeosphaeria (http://www.indexfungorum.org, Aug. 2015),
of which nine species have been transferred to Neo-
phaeosphaeria (Coniothyriaceae or Leptosphaeriaceae)or
Phaeosphaeriopsis (Phaeosphaeriaceae) based on ascospore
and conidial morphology, as well as sequence data from SSU
and ITS nrDNA (C^
amara et al. 2001, 2003), and 10 asexual
species with coniothyrium-like morphology have been added to
this genus based on molecular evidence (Trakunyingcharoen
et al. 2014, Verkley et al. 2014, Liu et al. 2015).
Paraphaeosphaeria michotii (Westend.) O.E. Erikss., Arkiv
Bot. 6: 405. 1967. Fig. 9.
Basionym:Sphaeria michotii Westend., Bull. Acad. R. Sci. Belg.,
Cl. Sci., s
er. 2, 7: 87. 1859.
In culture, both sexual and asexual morphs were observed. Asci
77–90 × 13–14.5 μm. Ascospores ellipsoid, 18–24 × 5–7μm
(av. 21.3 × 6.2 μm, n = 30), l/w 2.8–3.8 (av. 3.5, n = 30), with a
submedian primary septum (0.56–0.67; av. 0.61, n = 30), 2-
septate (1+1+0). Conidia broadly fusiform to ellipsoid,
8–14.5 × 4–7μm (av. 10.8 × 5.3 μm, n = 50), l/w 1.6–2.9 (av.
2.0, n = 50), echinulate.
Specimen examined:Japan, Tochigi, Utsunomiya, Ootanikannon, on dead
leaves of Typha latifolia, 29 Mar. 2007, K. Tanaka & Y. Harada, KT 2222 = HHUF
30142, culture MAFF 243861.
Notes: Morphological features of the sexual morph in our
material agree well with the description of P. m i c h o t i i
reported by several authors (Eriksson 1967, Shoemaker &
Eriksson 1967, Shoemaker & Babcock 1985, Ariyawansa
et al. 2014). The conidia we observed are somewhat larger
than those reported by Webster (1955;6–10 × 3–5μm) and
C^
amara et al. (2001;4–8×2.4–4.4 μm), but match those
reported by Sivanesan (1984;5–13.5 × 3 –5μm). A BLAST
search using ITS sequences showed 99.8 % similarity to
sequences from the ex-epitype of P. m i c h o t i i (GenBank
KJ939279; 519/520) deposited in GenBank by Ariyawansa
et al. (2014).
Spegazzinia Sacc., Michelia 2 (6): 37. 1880.
Type species:Spegazzinia ornata Sacc.
Notes: Based on the morphological features of basauxic con-
idiogenesis, Hyde et al. (1998) considered Spegazzinia to be a
possible member of the Apiosporaceae (Sordariomycetes),
although this was not fully resolved by molecular evidence (Hyde
et al. 2011, Crous & Groenewald 2013). We have assigned this
genus to the Didymosphaeriaceae (Fig. 1).
Spegazzinia deightonii (S. Hughes) Subram., J. Indian Bot.
Soc. 35: 78. 1956. Fig. 10.
Basionym:Spegazzinia tessarthra var. deightonii S. Hughes,
Mycol. Pap. 50: 65. 1953.
Specimens examined:Japan, Kagoshima, Isl. Yakushima, Kurio, on dead leaves
of Arundo donax, 14 Mar. 2007, K. Tanaka & H. Yonezawa, yone 66 = HHUF
E
DL
K
I
F
G
A
CHJ
B
Fig. 9. Paraphaeosphaeria michotii. A. Ascomata on the natural host surface; B. Ascoma in longitudinal section; C. Ascomatal wall; D. Ascus; E–H. Ascospores (arrowheads
indicate gelatinous sheath; H. in India ink); I. Conidioma in culture (on rice straw); J. Conidioma in longitudinal section; K, L. Conidia. A–C from KT 2222; D –L from culture KT
2222. Scale bars: A, I = 500 μm; B, J = 50 μm; C–H, K, L = 10 μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 93
30150, culture MAFF 243876; Okinawa, Isl. Iriomote, Inamori path, on dead
stems of herbaceous plant, 25 Sep. 2007, H. Yonezawa & K. Tanaka, yone
212 = HHUF 30151, culture MAFF 243877.
Note: The above collections match the original description and
illustration of S. tessarthra var. deightonii provided by Hughes
(1953b).
Spegazzinia tessarthra (Berk. & M.A. Curtis) Sacc., Syll. Fung. 4:
758. 1886. Fig. 11.
Basionym:Sporidesmium tessarthrum Berk. & M.A. Curtis, J.
Linn. Soc., Bot. 10: 355. 1868 (1869).
Specimen examined:Japan, Aomori, Towada, Sanbongi, Yagami, on balsa
wood, 9 Nov. 2002, S. Hatakeyama, SH 287 = HHUF 27691, culture MAFF
243875.
Notes: The features of the collection match the description of
S. tessarthra (Ellis 1971). The ITS sequences from the above
isolate were identical to those of S. tessarthra in GenBank
(GenBank JQ673429).
Lentitheciaceae Y. Zhang ter et al., Stud. Mycol. 64: 93. 2009.
Type genus:Lentithecium K.D. Hyde et al.
Katumotoa Kaz. Tanaka & Y. Harada, Mycoscience 46: 313.
2005.
Type species:Katumotoa bambusicola Kaz. Tanaka & Y. Harada.
Notes: Two bambusicolous genera, Katumotoa (Tanaka &
Harada 2005) and Neoophiosphaerella, formed a clade as re-
ported in previous molecular studies (Schoch et al. 2009, Tanaka
et al. 2009, Zhang et al. 2012), but these have ascospores with
distinct features, i.e., apiosporous in Katumotoa and multi-
septate scolecosporous in Neoophiosphaerella. Furthermore,
sequence similarity between the type species of both genera was
82.7 % (417/504) in their ITS regions, suggesting that they are
not congeneric. Although a correlation between phylogenetic
relationships and host preferences has been noted in these
bambusicolous fungi (Zhang et al. 2012), discovery of additional
species in both monotypic genera will be needed to confirm this.
Katumotoa bambusicola Kaz. Tanaka & Y. Harada, Myco-
science 46: 313. 2005. Fig. 12.
Specimen examined:Japan, Iwate, Nishine, Hirakasa, near Yakebashiri, Mt.
Iwate, on Sasa kurilensis, 19 Oct. 2003, K. Tanaka (paratype KT 1517a = HHUF
28661, culture ex-paratype JCM 13131 = MAFF 239641).
Keissleriella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl.,
Abt. 1128: 582. 1919.
Type species:Keissleriella aesculi (Höhn.) Höhn.
Notes:Keissleriella is characterised by an ostiolar neck covered
by short dark setae. Munk (1953) introduced Trichome-
tasphaeria, which can be separated from Keissleriella by their
host preferences and the morphological features of hamathe-
cium and ascospores. Trichometasphaeria occurs on herba-
ceous plants and has cellular pseudoparaphyses and septate
ascospores, while Keissleriella occurs on woody plants and has
trabecular pseudoparaphyses and 1-septate ascospores (Munk
1957, Barr 1990a, 1992). However, Bose (1961) treated Tri-
chometasphaeria as a synonym of Keissleriella, and this opinion
was accepted by several authors (e.g., Eriksson 1967, Dennis
1978, Sivanesan 1984). We also regard these two genera as
B
A
CDE
Fig. 10. Spegazzinia deightonii. A, B. Sporodochia on the natural host surface; C, D. Conidiophores and conidiogenous cells (arrowheads indicate basauxic conidiogenesis); E.
Conidia (arrowheads indicate atype conidia, arrows indicate btype conidia). A from yone 66; B–E from culture yone 66. Scale bars: A, B = 500 μm; C–E=10μm.
D
C
BA
Fig. 11. Spegazzinia tessarthra. A. Sporodochia on the natural host surface; B, C. Conidiophores and conidiogenous cells (arrowheads indicate basauxic conidiogenesis); D.
Conidia (arrowheads indicate atype conidia, arrows indicate btype conidia). All from SH 287. Scale bars: A = 500 μm; B –D=10μm.
TANAKA ET AL.
94
congeneric as species with the features of Trichometasphaeria
(e.g., K. gloeospora, the type species of Trichometasphaeria)
and Keissleriella (e.g., K. cladophila;Corbaz 1956, Bose 1961)
form a clade (Fig. 1), although the type of the latter genus
(K. sambucina) is not included in our analysis.
Keissleriella breviasca Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811304. Fig. 13.
Etymology: From the Latin brevi- meaning short, in reference to
the ascus length.
Ascomata scattered, erumpent, 95–115 μm high, 130 –165 μm
diam. Ostiolar neck papillate, 18–23(–50) μm long, covered with
1-celled, thick-walled, dark brown to almost black setae. Asco-
matal wall 7.5–11.5 μm thick at sides, composed of 3–4 layers
of polygonal to subglobose cells of 2.5–10 × 2.5–5μm. Pseu-
doparaphyses cellular, septate, branched and anastomosed,
2–3.5 μm wide. Asci fissitunicate, clavate to cylindrical,
51–66 × 7–11 μm (av. 59.0 × 8.7 μm, n = 39), rounded at the
apex and with a shallow ocular chamber, short-stalked (5–12 μm
long), with 4 biseriate ascospores. Ascospores narrowly fusiform,
17.5–24.5 × 3.5–5μm (av. 21.1 × 4.3 μm, n = 64), l/w (3.7–)
4.0–5.7(–6.3) (av. 4.9, n = 64), with a nearly median primary
septum (0.44–0.53; av. 0.49, n = 48), 3-septate (1+1+1), slightly
constricted at the septa, hyaline, smooth, with an entire sheath;
sheath gelatinous and 8–11 μm wide when fresh, delimited and
1–2μm wide when dry.
Colonies on PDA (after 4 wk) attaining a diam of 1.0–1.3 cm,
coral with white margin; reverse red to flesh; sienna pigment
produced. In culture coelomycetous asexual morph formed.
Conidiomata pycnidial, 70–90 μm high, 120–200 μm diam,
subglobose to hemispherical in section. Conidiomatal wall
7–15 μm thick at sides, composed of thin-walled, flattened cells.
Conidiogenous cells cylindrical to doliiform, 8–13 × 2.5–3.5 μm,
holoblastic. Conidia cylindrical to bone-shaped, 0–3-septate,
hyaline, smooth, 11–20 × 3–4μm (av. 15.2 × 3.7 μm, n = 20), l/w
3.7–5.0 (av. 4.1, n = 20), without sheath. Sometimes, sexual
morph formed. Ascospores 19–23 × 3.5–4.5 μm.
DEPON
ML
C
AB
FHIJK
G
Fig. 13. Keissleriella breviasca. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Apical setae of ascoma; E. Ascomatal wall; F. Pseudo-
paraphyses; G. Ascus; H–K. Ascospores (arrowheads indicate gelatinous sheath); L, M. Ascomata and conidiomata in culture (on rice straw); N. Ascoma produced in culture;
O. Conidioma produced in culture; P. Conidia. A–C, E, I from KT 649; D, G, H from KT 540; F, J from KT 581; K from culture KT 581; L–O from culture KT 649; P from culture
KT 540. Scale bars: A, L = 2 mm; B, M = 500 μm; C, N, O = 20 μm; D–K, P = 10 μm.
B F
A
DG
E
C
Fig. 12. Katumotoa bambusicola. A. Ascomata on the natural host surface; B, C. Ascomata in longitudinal section; D. Ascomatal wall (in lactophenol cotton blue); E.
Pseudoparaphyses; F. Ascus; G. Ascospore (arrowheads indicate gelatinous sheath). A–D from KT 1517; E–G from culture KT 1517. Scale bars: A = 1 mm; B, C = 100 μm;
D–G=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 95
Specimens examined:Japan, Aomori, Hirosaki, Campus of Hirosaki Univ., on
dead culms of Dactylis glomerata, 13 Jun. 2001, K. Tanaka, KT 540 = HHUF
27715, culture JCM 19413 = MAFF 239476; ibid., 20 Jul. 2001, KT 581 = HHUF
27717, culture JCM 19414 = MAFF 243843; Aomori, Hirakawa, Hirakawa-river,
riverbank, on dead culms of Dactylis glomerata, 5 Aug. 2001, K. Tanaka (ho-
lotype KT 649 = HHUF 27718, culture ex-type CBS 139691 = JCM
19415 = MAFF 243844).
Notes: To date, 50 species have been described as Keissleriella
(or Trichometasphaeria; MycoBank, http://www.mycobank.org,
April 2015), but K. breviasca can be distinguished from all
known species by its short asci with consistently four asco-
spores (Fig. 13G). The asexual morph of K. breviasca is similar
to that of K. gallica reported as “Ascochyta sp.”(Sivanesan
1984), but the conidia of K. breviasca have 0–3septa(vs.
0–1septainK. gallica). A collection of K. culmifida with 4-
spored asci has sometimes been reported (Dennis 1978,
Ridley 1988), which is most probably conspecificwith
K. breviasca.
Keissleriella culmifida (P. Karst.) S.K. Bose, Phytopath. Z. 41:
188. 1961. Fig. 14.
Basionym:Leptosphaeria culmifida P. Karst., Bidr. K€
ann. Finl.
Nat. Folk 23: 103. 1873.
Ascomata 120–145 μm high, 120–150 μm diam, with dark
brown setae of 30–58 × 3–5μm around the ostiole. Asci 8-
spored, 70.5–84 × 10.5–13 μm. Ascospores fusiform,
17.5–22 × 4–5.5 μm (av. 19.4 × 4.9 μm, n = 36), l/w 3.5–5.0 (av.
3.9, n = 36), with nearly median primary septum (0.47–0.53; av.
0.51, n = 35), 3-septate (1+1+1), surrounded by an entire
gelatinous sheath, 3–9μm wide.
In culture sexual morph formed. Asci 90–127.5 × 10.5–15 μm
(av. 102.9 × 13.0 μm, n = 40). Ascospores fusiform,
19–26 × 5–7μm (av. 22.5 × 5.9 μm, n = 60), l/w 3.3–4.5 (av. 3.8,
n = 60), with a nearly median primary septum (0.46–0.55; av.
0.50, n = 60). Asexual morph not observed.
Specimens examined:Japan, Hokkaido, Isl. Rishiri, Forest park, on dead stems
of Agrostis flaccida, 29 Jul. 2007, K. Tanaka & G. Sato, KT 2308 = HHUF 30135,
culture JCM 19416 = MAFF 243848; Iwate, Hachimantai, Top of Aspite line, on
dead leaves of Festuca sp. 25 Jul. 2009, K. Tanaka & Y. Harada, KT
2642 = HHUF 30136, culture JCM 19417 = MAFF 243849.
Notes: We identified these specimens as K. culmifida, based on
the description (Karsten 1873, Holm 1957, Bose 1961) and
illustration (Berlese 1894, Eriksson 1967) of this species. How-
ever, ITS sequences from the above materials completely
matched with those from the ex-type of K. poagena (GenBank
KJ869112), a species recently published by Crous et al. (2014a).
Keissleriella poagena is morphologically close to K. culmifida,
and thus taxonomic reassessment of these two species will be
needed.
Keissleriella gloeospora (Berk. & Curr.) S.K. Bose, Phytopath.
Z. 41: 190. 1961. Fig. 15.
Basionym:Sphaeria gloeospora Berk. & Curr., Ann. Mag. Nat.
Hist., Ser. 3 7: 454. 1861.
Ascomata 130–230 μm high, 350–480 μm diam, with dark
brown setae (up to 60 μm long) around the ostiole. Asci clavate,
80–118 × 14–19 μm (av. 97.2 × 16.8 μm, n = 20), short-stalked
(10–25 μm long). Ascospores fusiform to clavate,
21.5–33 × 6–9μm (av. 28.0 × 7.3 μm, n = 50), l/w 3.3–4.9 (av.
3.8, n = 50), with a supramedian primary septum (0.39–0.48; av.
0.43, n = 50), 4–6-septate (1+1+2, 1+1+3, 2+1+2, 2+1+3),
sometimes with a vertical septum in central cells, surrounded by
an entire sheath (6–10 μm wide when fresh, 2–3μm wide when
dry).
In culture spermatial morph formed. Spermatia
3.3–4.9 × 1.2 –1.8 μm(av.4.1×1.5μm, n = 20), hyaline, cylindrical.
Specimen examined:Japan, Aomori, Hirosaki, Campus of Hirosaki Univ., on
dead culms of Setaria faberii, 4 Nov. 2001, K. Tanaka, KT 829 = HHUF 27704,
culture MAFF 239474.
Note: The above material matches well the descriptions of
K. gloeospora provided by Bose (1961) and Shearer et al. (1993).
Keissleriella quadriseptata Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811305. Fig. 16.
C
AB
EG
D
K
J
I
H
F
Fig. 14. Keissleriella culmifida. A. Ascomata on the natural host surface; B. Ascomata in culture (on rice straw); C. Ascoma in longitudinal section; D. Apical setae of ascoma;
E. Ascomatal wall; F. Pseudoparaphyses; G. Ascus; H–J. Ascospores (arrowheads indicate gelatinous sheath); K. Germinating ascospore. A, C, E, K from KT 2642; B, D, F, G,
I, J from culture KT 2308; H from KT 2308. Scale bars: A, B = 500 μm; C, D, K = 20 μm; E–J=10μm.
TANAKA ET AL.
96
Etymology: Referring to the 4-septate ascospores.
Ascomata scattered to 2–4 grouped, immersed to erumpent,
globose in section, 210–320 μm high, 170–310 μm diam.
Ostiolar neck central, papillate, 35–50 μm long, 55–62 μm diam,
with setae; setae bluntly pointed, aseptate, slightly waved, dark
brown to black, 37–93 μm long, 2.5–4μm wide at the base.
Ascomatal wall in longitudinal section uniformly 15–20 μm thick,
composed of 5–6 layers of polygonal, slightly thick-walled,
4.5–16.5 × 3–7.5 μm, brown cells. Pseudoparaphyses cellular,
septate, branched and anastomosed, 2–2.5 μm wide. Asci fis-
situnicate, clavate, 78.5–107.5 × 12–14.5 μm (av.
92.8 × 13.1 μm, n = 30), rounded at the apex and with a shallow
ocular chamber, short-stalked (5–24 μm long; av. 13.1 μm,
n = 26), with 8 biseriate ascospores. Ascospores clavate,
19–24.5 × 5–7μm (av. 21.5 × 5.8 μm, n = 50), l/w 3.3–4.4 (av.
3.7, n = 50), with a supramedian primary septum (0.38–0.45; av.
0.42, n = 50), 4-septate (1+1+2), slightly constricted at the
G
FE
B
A
K
D
C
HI
J
Fig. 15. Keissleriella gloeospora. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Apical setae of ascoma; E. Ascomatal wall; F. Pseu-
doparaphyses; G. Ascus; H–J. Ascospores (arrowheads indicate gelatinous sheath); K. Spermatia. A–J from KT 829; K from culture KT 829. Scale bars: A = 1 mm;
B = 500 μm; C = 50 μm; D–K=10μm.
G
D
C
AB
F
MPQ
E
K
J
I
H
ON
L
Fig. 16. Keissleriella quadriseptata. A, B. Ascomata on the natural host surface; C, D. Ascomata in longitudinal section; E. Apical setae of ascoma; F. Ascomatal wall; G.
Pseudoparaphyses; H. Ascus; I–K. Ascospores (arrowheads indicate gelatinous sheath); L. Germinating ascospore; M. Conidiomata in culture (on rice straw); N. Conidioma in
longitudinal section; O. Conidiomatal wall; P. Conidium; Q. Senescent conidium. A –L from KT 2292; M–Q from culture KT 2292. Scale bars: A, M = 500 μm; B = 200 μm; C, D,
N=50μm; E–L, O–Q=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 97
primary septum, hyaline, smooth, with an entire gelatinous
sheath of 4–6μm wide.
Colonies on PDA (after 2 wk) attaining a diam of 1.7–2.2 cm,
pale luteous; reverse rust; no pigment produced. In culture
asexual morph formed. Conidiomata pycnidial, 200–380 μm
high, 160–310 μm diam, globose in section. Ostiolar neck
papillate, 60–90 μm long, 50–75 μm wide. Conidiomatal wall
13–25 μm thick, composed of 4–6 layers of polygonal cells
(5–17 × 2.5–6.5 μm). Conidiophores absent. Conidiogenous
cells lageniform, holoblastic. Conidia cylindrical, rounded at the
apex, slightly truncate at the base, straight, aseptate, hyaline,
smooth, 25–32 × 6–8.5 μm (av. 28.4 × 7.2 μm, n = 30), l/w
3.3–4.8 (av. 3.9, n = 30), without sheath; senescent spores
brown, 0–1-septate.
Specimen examined:Japan, Hokkaido, Isl. Rishiri, Beach near Ryuzinnoiwa, on
dead culms of Dactylis glomerata, 28 Jul. 2007, K. Tanaka & G. Sato (holotype KT
2292 = HHUF 30137, culture ex-type CBS 139692 = JCM 19418 = MAFF 243850).
Notes: This species is characterised by its consistently 4-septate
ascospores (1+1+2). Phylogenetically, it is closest to
K. gloeospora (97.8 %; 904/924 in tef1), but the latter has larger
ascospores (21.5–33 × 6–9μm) with 4–6 septa (1+1+2, 1+1+3,
2+1+2, 2+1+3). In culture, K. quadriseptata produced an asexual
morph with cylindrical, aseptate, hyaline conidia, but only a
spermatial morph was found in K. gloeospora.
Keissleriella taminensis (H. Wegelin) S.K. Bose, Phytopath. Z.
41: 190. 1961. Fig. 17.
Basionym:Leptosphaeria taminensis H. Wegelin, Mitt. Thürgau.
Naturf. Ges. 12: 173. 1896.
Ascomata 190–280 μm high, 180–300 μm diam. Asci clavate,
70–106 × 11.5–16 μm (av. 84.3 × 13.3 μm, n = 53), short-stalked
(8–25 μm long; av. 15.5 μm, n = 27). Ascospores fusiform,
19–25 × 4–7μm (av. 21.9 × 5.9 μm, n = 103), l/w 3.0–4.5(–5.0)
(av. 3.7, n = 103), with a supramedian primary septum
(0.43–0.51; av. 0.47, n = 100), 4–5-septate (1+1+2, 2+1+2),
surrounded by an entire sheath (3–10 μm wide when fresh,
1–2μm wide when dry).
In culture spermatial morph formed. Spermatia cylindrical, hya-
line, 3.5–8.8 × 1.3–2.3 μm (av. 5.5 × 1.8 μm, n = 60). Sometimes
sexual morph observed.
Specimens examined:Japan, Aomori, Hirosaki, Sanpinai, on dead stems of
herbaceous plant, 17 Jul. 2001, K. Tanaka, KT 571 = HHUF27707, culture MAFF
243846; Aomori, Hirosaki, Kadoke, Oowasawa-river, riverbank, on dead stems of
herbaceous plant, 29 Jul. 2001, K. Tanaka, KT 594 = HHUF27709, culture MAFF
243847; ibid., 14 Aug. 2001, K. Tanaka, KT 678 = HHUF 27711, culture MAFF
239475.
Note: These materials agree well with the description of
K. taminensis except for the slightly wider ascospores in our
collections (4–7μmvs. 4.5–5μm; Bose 1961).
Keissleriella yonaguniensis Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811307. Fig. 18.
Etymology: Referring to the collection site.
Ascomata scattered, immersed to erumpent, globose in section,
100–170 μm high, 100–180 μm diam. Ostiolar neck central,
papillate to terete, 37–45 μm long, 32–35 μm diam, with setae;
setae bluntly pointed, aseptate, straight, dark brown to black,
20–30 μm long, 3–4.5 μm wide at the base. Ascomatal wall in
longitudinal section uniformly 7.5–10 μm thick, composed of 3–5
layers of polygonal, thin-walled, flattened, 3.5–7.5 × 1.5–2.5 μm,
brown cells. Pseudoparaphyses cellular, septate, branched and
anastomosed, 1.5–2.5 μm wide (3–4.5 μm wide at below). Asci
fissitunicate, cylindrical, 65–99.5(–112.5) × 10.5–14.5 μm (av.
80.1 × 12.1 μm, n = 35), rounded at apex and with a shallow
ocular chamber, with a short stipe of 5–13 μm long, 8-spored.
Ascospores cylindrical with rounded ends, 15–20 × 4.5–6.5 μm
(av. 18.1 × 5.3 μm, n = 50), l/w 3.0–3.9 (av. 3.5, n = 50), with a
nearly median primary septum (0.47–0.53; av. 0.50, n = 50), 5-
septate (2+1+2), yellow, smooth, with an entire gelatinous sheath
of 3–5μm wide (later diffuse up to 10 μm wide).
Colonies on PDA (after 4 wk) attaining a diam of 5.2–5.7 cm,
white to pale luteous; reverse similar; no pigment produced. In
culture sexual morph identical to that formed on the natural host
produced.
H
E
C
A
GJK
B
F
DI
Fig. 17. Keissleriella taminensis. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Apical setae of ascoma; E. Ascomatal wall; F. Pseu-
doparaphyses; G. Ascus; H–J. Ascospores (arrowheads indicate gelatinous sheath); K. Spermatia. A –E, G, I from KT 594; F, H from KT 571; J from KT 678; K from culture KT
594. Scale bars: A = 2 mm; B = 500 μm; C = 50 μm; D–K=10μm.
TANAKA ET AL.
98
Specimen examined:Japan, Okinawa, Isl. Yonaguni, Kubura pond, on dead
leaves of Typha latifolia, 23 Nov. 2008, K. Tanaka & K. Hirayama (holotype KT
2604 = HHUF 30138, culture ex-type CBS 139693 = JCM 19419 = MAFF 243851).
Notes: Among the 50 described species in Keissleriella (or Tri-
chometasphaeria), six are known to have 5-septate ascospores like
K. yonaguniensis. These are K. abruptipapilla,K. gloeospora,
T. papillisetosa,K. pindaundeensis,T. populi and K. taminensis.
Among these, K. gloeospora,T. populi and K. taminensis differ from
K. yonaguniensis in having asymmetrically-septate ascospores
(Bose 1961, Barr 1992). Keissleriella pindaundeensis (Kobayasi
1971)andT. papillisetosa (Yuan & Barr 1994) are distinguished
from K. yonaguniensis by larger ascospores that are more than
20 μmlong.Keissleriella abruptipapilla (Barr 1990a) is similar but its
ascospores are verruculose. In our phylogenetic tree (Fig. 1),
K. yonaguniensis grouped with K. linearis (= Lentithecium lineare;
Zhang et al. 2009c), Murilentithecium with muriform ascospores
and conidia (Wanasinghe et al. 2014), and Phragmocamar-
osporium spp. with phragmosporous conidia (Wijayawardene et al.
2015), although this clade received no support. We include this
species as Keissleriella based on the morphology of its sexual
morph pending further studies of related taxa.
Lentithecium K.D. Hyde et al., Fungal Divers. 38: 234. 2009.
Type species:Lentithecium fluviatile (Aptroot & Van Ryck.) K.D.
Hyde et al.
Notes:Lentithecium was established by Zhang et al. (2009c)
using L. fluviatile as the type species. This genus was charac-
terised by the lenticular ascomata, but reexamination based on
the holotype of L. fluviatile revealed that the species has globose
ascomata (Hyde et al. 2013).
Lentithecium clioninum (Kaz. Tanaka et al.) Kaz. Tanaka & K.
Hiray., comb. nov. MycoBank MB811308. Fig. 19.
Basionym:Massarina clionina Kaz. Tanaka et al., Mycoscience
46: 288. 2005.
Ascomata 210–280 μm high, 330–430 μm diam. Ostiolar neck
50–75 μm long, 75–125 μm wide. Ascomatal wall 15–23 μm
thick at sides. Pseudoparaphyses 2μm wide. Asci clavate,
(81.5–)86–118(–128) × 15–19(–21) μm (av. 100.1 × 17.2 μm,
n = 70), short-stalked (5–23 μm long). Ascospores fusiform,
(26–)27.5–34.5(–37) × 7–10(–11) μm (av. 31.0 × 8.7 μm,
n = 70), l/w 3.2–4.1 (av. 3.6, n = 70), with a septum mostly
median (0.48–0.52; av. 0.50, n = 64), with a wing-like sheath
staining with Black-Blue ink when fresh.
In culture ascomatal morph formed. Ascospores
27–35 × 8–11 μm (av. 31.0 × 9.3 μm, n = 50). No asexual morph
observed.
Specimens examined:Japan, Hokkaido, Akkeshi, Bekanbeushi-river (near
Bekanbeushi bridge), on submerged twigs of woody plant, 2 Jun. 2003, K. Tanaka
& S. Hatakeyama (holotype KT 1149A = HHUF 28199, culture ex-type CBS
139694 = JCM 12703 = MAFF 239293); Hokkaido, Akkeshi, Toraibetsu-river, on
submerged twigs of woody plant, 3 Jun. 2003, K. Tanaka & S. Hatakeyama
(paratype KT 1220 = HHUF 28213, culture ex-paratype MAFF 243839).
Notes: This species was previously described as Massarina
(Tanaka et al. 2005b), but we here transfer it to Lentithecium,
because it has morphological similarities with L. fluviatile, the
type species of the genus (see Fig. 43 in Zhang et al. 2012). Both
have globose ascomata composed of small polygonal peridial
cells, short pedicellate asci, and fusiform ascospores with obtuse
ends. In our phylogenetic tree (Fig. 1), this species and
L. pseudoclioninum nested on a well-supported branch (100 %)
with L. fluviatile.
Lentithecium pseudoclioninum Kaz. Tanaka & K. Hiray., sp.
nov. MycoBank MB811309. Fig. 20.
Etymology: Referring to its resemblance to Lentithecium
clioninum.
Ascomata scattered to 2–3 grouped, immersed to erumpent,
globose in section, 200–220 μm diam, with sparse brown hy-
phae around ascomata. Ostiolar neck central, papillate,
40–50 μm long, 30 μm diam, with periphyses. Ascomatal wall in
longitudinal section uniformly 17.5–20 μm thick, composed of
5–6 layers of polygonal to subglobose, 5–12.5 × 2 μm, brown
cells. Pseudoparaphyses septate, branched and anastomosed,
B
FK
J
A
C
D
EG
HI
Fig. 18. Keissleriella yonaguniensis. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Apical setae of ascoma; E. Ascomatal wall; F.
Pseudoparaphyses; G. Asci; H–K. Ascospores (arrowheads indicate gelatinous sheath; K. in India ink). All from KT 2604. Scale bars: A = 2 mm; B = 500 μm; C = 50 μm;
D–K=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 99
2–3μm wide. Asci fissitunicate, clavate, 62.5–116 × 14 –25 μm
(av. 92.0 × 18.4 μm, n = 95), rounded at the apex and with a
shallow ocular chamber, short-stalked (5–17.5 μm long; av.
9.9 μm, n = 67), with 8 biseriate ascospores. Ascospores clavate
to broadly fusiform, slightly curved, 22–39 × 6.5–11.5 μm (av.
29.2 × 8.5 μm, n = 77), l/w (2.4–)2.8–4.1(–4.6) (av. 3.5, n = 77),
with a supramedian septum (0.43–0.50; av. 0.48, n = 73),
constricted at the septum, hyaline, smooth, with an amorphous
gelatinous sheath (1–4μm wide) staining with Black-Blue ink
when in fresh condition.
Colonies on PDA (after 4 wk) attaining a diam of 2.7–3.0 cm,
smoke grey; reverse vinaceous buff to citrine; no pigment pro-
duced. In culture sexual morph identical to that formed on the
natural host produced.
Specimens examined:Japan, Aomori, Hirosaki, Aoki, Mohei pond, on sub-
merged twigs of woody plant, 3 May 2003, K. Tanaka & N. Asama, KT
1111 = HHUF 29053, culture JCM 19421 = MAFF 243840; ibid. (holotype KT
1113 = HHUF 29055, culture ex-type CBS 139695 = JCM 19422 = MAFF
243841).
Notes: Morphologically, this species is close to L. clioninum, but
is clearly separated from it on the basis of its smaller ascospores
with a supramedian septum. The wing-like sheath of ascospores
found in L. clioninum was not observed in L. pseudoclioninum.
Neoophiosphaerella Kaz. Tanaka & K. Hiray., gen. nov.
MycoBank MB811310.
Etymology: After its morphological similarity to Ophiosphaerella.
Ascomata scattered to grouped, erumpent, subglobose to
hemispherical with flattened base. Ostiolar neck central, terete,
flush, covered by black clypeus. Ascomatal wall at sides
composed of several layers of thin-walled brown cells. Pseu-
doparaphyses numerous, cellular, branched. Asci fissitunicate,
cylindrical, short-stalked, with 8 parallel or twisted ascospores.
Ascospores filiform, multiseptate, hyaline to pale yellowish
brown, smooth, surrounded by a sheath. Asexual morph
unknown.
Type species:Neoophiosphaerella sasicola (Nagas. & Y. Otani)
Kaz. Tanaka & K. Hiray.
Notes:Neoophiosphaerella sasicola, the type species of this
genus, was originally described as a species of Phaeosphaeria
(Nagasawa & Otani 1977) and later transferred to Ophios-
phaerella (Shoemaker & Babcock 1989). These two genera,
however, belong to the Phaeosphaeriaceae in the Pleosporineae
(C^
amara et al. 2000, Schoch et al. 2009, Phookamsak et al.
2014), a family distantly related to the Lentitheciaceae.We
therefore propose a new genus, Neoophiosphaerella,to
accommodate this species. Ophiosphaerella has globose to
subglobose ascomata with a papillate ostiolar neck
(Phookamsak et al. 2014), while Neoophiosphaerella is char-
acterised by hemispherical ascomata without papilla but being
covered by clypei.
B
A
CED
H
JF I
G
Fig. 20. Lentithecium pseudoclioninum. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G–I.
Ascospores (arrowheads indicate gelatinous sheath; all in Black-Blue ink); J. Germinating ascospore. All from KT 1113.Scale bars: A = 1 mm; B = 500 μm; C = 50 μm; D –J=10μm.
E
H
CD
A
FGI
B
Fig. 19. Lentithecium clioninum. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G–I.
Ascospores (arrowheads indicate gelatinous sheath; H, I. in Black-Blue ink). A–D from KT 1149A; E–G from KT 1220; H, I from culture KT 1149A. Scale bars: A = 1 mm;
B = 500 μm; C = 50 μm; D–I=10μm.
TANAKA ET AL.
100
Neoophiosphaerella sasicola (Nagas. & Y. Otani) Kaz. Tanaka
& K. Hiray., comb. nov. MycoBank MB811312. Fig. 21.
Basionym:Phaeosphaeria sasicola Nagas. & Y. Otani, Rep.
Tottori Mycol. Inst. 15: 39. 1977.
Specimens examined:Japan, Hokkaido, Ebestu, Nopporo, on Sasa senanensis,
15 May 1972, E. Nagasawa (holotype of Phaeosphaeria sasicola TMI 3176);
Hokkaido, Isl. Rebun, Funadomari, Akaiwa, on dead culms of Sasa kurilensis,5
Jun. 2004, K. Tanaka (epitype designated here KT 1706 = HHUF 29443,
MBT202864, culture ex-epitype CBS 120247 = JCM 13134 = MAFF 239644).
Note: The collection HHUF 29443 is designated as epitype for
N. sasicola, the type species of Neoophiosphaerella.
Setoseptoria Quaedvl. et al., Stud. Mycol. 75: 382, 2013.
Type species:Setoseptoria phragmitis Quaedvl. et al.
Notes:Quaedvlieg et al. (2013) established the genus Seto-
septoria typified by S. phragmitis on Phragmites. The sexual
morph of this coelomycetous genus is presently unknown.
Setoseptoria arundinacea (Sowerby) Kaz. Tanaka & K. Hiray.,
comb. nov. MycoBank MB811313. Fig. 22.
Basionym:Sphaeria arundinacea Sowerby, Col. Fig. Engl. Fung.
3: 139, t. 336. 1803.
Specimens examined:Japan, Aomori, Hirosaki, Sanpinai, on dead culms of
Phragmites australis, 1 Jul. 2001, K. Tanaka, KT 552 = HHUF 27543, culture
MAFF 239460; Aomori, Hirosaki, Kadoke, Oowasawa-river, on dead culms of
Phragmites australis, 29 Jul. 2001, K. Tanaka, KT 600 = HHUF 27544, culture
MAFF 243842.
Notes: This species has been placed in various pleosporalean
genera, such as Leptosphaeria (see Crane & Shearer 1991),
Lophiostoma (Hyde et al. 2002), Massarina (Leuchtmann 1984),
Metasphaeria (Vasilyeva 1998), and Phaeosphaeria
(Hedjaroude 1968). Most recently, it has been transferred to
Lentithecium based on the results of phylogenetic analyses
using SSU + LSU nrDNA and rpb2 (Zhang et al. 2009c).
However, later molecular studies (Schoch et al. 2009, Shearer
et al. 2009, Zhang et al. 2009b, 2012, Liu et al. 2011), as well
as our own work (Fig. 1), do not support this placement. This
species and its phenotypically and phylogenetically close rela-
tive Massarina magniarundinacea (Tanaka et al. 2004) do not
belong to any genera previously suggested. They should,
therefore, be transferred to another genus. One candidate
genus to accommodate these species would be Setoseptoria.
The monotypic genus Setoseptoria typified by S. phragmitis
was introduced for a stagonospora-like pycnidial coelomycete
with (1–)3-septate, subcylindrical, hyaline conidia (Quaedvlieg
et al. 2013). The sexual morph of Setoseptoria is unknown.
In contrast, two massarina-like species, M. arundinacea and
M. magniarundinacea, have been reported to produce only
sexual morphs in culture (Lucas 1968, Leuchtmann 1984,
Tanaka & Harada 2003b, Tanaka et al. 2004). Although there
is no example of production of stagonospora-like asexual
morphs from the massarina-like sexual morphs in culture,
congeneric relationships have been suggested several times
between Stagonospora elegans (Aptroot 1998, Eriksson &
Hawksworth 2003)orS. vexata (Grove 1935) and
M. arundinacea, based on their close association on the same
host tissue. Furthermore, two stagonospora-like species
(Setoseptoria phragmitis and “Stagonospora”macropycnidia)
and two massarina-like species (M. arundinacea and
M. magniarundinacea) form a strongly supported clade (Fig. 1)
and sequence similarities between these species in the LSU
region are considerably higher (826/834 = 99.0 %). We there-
fore tentatively assign the two massarina-like species to the
genus Setosphaeria, although asexual morphs of these species
are presently unknown. A species with both sexual and asexual
morphs will be required to confirm the validity of our generic
treatment.
The two isolates of S. arundinacea that we examined did not
form a clade with S. arundinacea from GenBank (Fig. 1). This
species has been reported many times as a common species on
Phragmites culms (Aptroot 1998, Tanaka & Harada 2003b), but
S. arundinacea may consist of several cryptic species with close
morphological resemblance.
AB
G
F
CED
Fig. 21. Neoophiosphaerella sasicola. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G.
Ascospore (arrowheads indicate gelatinous sheath; in India ink). All from KT 1706. Scale bars: A = 1 mm; B = 500 μm; C = 100 μm; D–G=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 101
Setoseptoria magniarundinacea (Kaz. Tanaka & Y. Harada)
Kaz. Tanaka & K. Hiray., comb. nov. MycoBank MB811314.
Fig. 23.
Basionym:Massarina magniarundinacea Kaz. Tanaka & Y.
Harada, Mycotaxon 90: 349. 2004.
Ascomata subglobose to hemispherical with flattened base,
150–280 μm high, 310–410 μm diam. Ostiolar neck central,
short papillate. Ascomatal wall 12.5–20 μm thick at sides.
Pseudoparaphyses cellular, 1.5–3.5 μm wide. Asci (119–)
125–182.5(–200) × 25–35(–47.5) μm (av.151.5 × 30.2 μm,
n = 37). Ascospores 67–82 × 6.5–9μm (av. 74.0 × 7.7, n = 50), l/
w 8.4–11.0 (av. 9.6, n = 50), with a submedian primary septum
(0.52–0.57; av. 0.54, n = 50).
In culture the ascomatal morph is similar to that observed on the
natural host.
Specimen examined:Japan, Hokkaido, Akkeshi, Ariake, small stream, on sub-
merged stems of herbaceous plant, 3 Jun. 2003, K. Tanaka & S. Hatakeyama
(holotype KT 1174 = HHUF 28293, culture ex-type CBS 139702 = MAFF
239294).
Note: The ascospores of S. magniarundinacea are most similar
in shape and colour to those of S. arundinacea, but are
considerably larger (67–82 × 6.5–9μmvs.23–40 × 3.5–6μm,
Tanaka et al. 2004).
Tingoldiago K. Hiray. & Kaz. Tanaka, Mycologia 102: 740. 2010.
Type species:Tingoldiago graminicola K. Hiray. & Kaz. Tanaka.
Notes:Tingoldiago, found in freshwater environments, is
characterised by lens-shaped ascomata and narrowly fusiform
ascospores, each of which has an elongated sheath
(Hirayama et al. 2010;Fig. 27). This genus, however, was
regarded as a synonym of Lentithecium, despite the fact that
the Lentithecium clade that included Tingoldiago was not well-
supported (21 %; Zhang et al. 2012). When Lentithecium was
established, the lenticular ascomata with simple peridial
structure and hyaline 1-septate ascospores were emphasised
as important characters to define the genus (Zhang et al.
2009c), but this generic circumscription is incorrect (see
Notes in Lentithecium).
Tingoldiago graminicola K. Hiray. & Kaz. Tanaka, Mycologia
102: 740. 2010. Fig. 24.
Specimens examined:Japan, Hokkaido, Isl. Rishiri, Himenuma (pond), on
submerged culms of Phragmites australis, 27 Jul. 2007, K. Hirayama & K.
Tanaka (holotype KH 68 = HHUF 30009, culture ex-type JCM 16485 = NBRC
106131); ibid., 25 Jul. 2008, K. Hirayama & K. Tanaka (paratype KH
155 = HHUF 30010, culture ex-paratype JCM 16486 = NBRC 106132); Aomori,
Hirosaki, Kadoke, Oowasawa-river, on submerged culms of Phragmites
japonica, 28 Sep. 2002, K. Tanaka (paratype KT 891 = HHUF 27882, culture
ex-paratype MAFF 239472).
Notes: The morphological features of this species have been
described by Hirayama et al. (2010).Tingoldiago graminicola
and Setoseptoria arundinacea, formerly treated as Lentithecium
by Zhang et al. (2009c, 2012), have lens-shaped ascomata, but
are not in the clade of Lentithecium s. str. (Fig. 1). Instead,
Lentithecium is restricted to species with globose ascomata (Figs
19C, 20C).
E
D
A
B H
FC G
Fig. 22. Setoseptoria arundinacea. A. Surface view of ascomata; B, C. Ascomata in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G, H. Ascospores
(arrowheads indicate gelatinous sheath). All from KT 600. Scale bars: A = 500 μm; B = 100 μm; C = 50 μm; D–H=10μm.
I
G
F
E
C
B
A
H
D
Fig. 23. Setoseptoria magniarundinacea. A, B. Surface view of ascomata; C. Ascoma in longitudinal section (in lactophenol cotton blue); D. Ascomatal wall (in lactophenol
cotton blue); E. Pseudoparaphyses; F. Ascus; G, H. Ascospores (arrowheads indicate gelatinous sheath); I. Germinating ascospore. A, B, E–H from culture KT 1174; C, D, I
from KT 1174. Scale bars: A = 1 mm; B = 500 μm; C = 50 μm; D–I=10μm.
TANAKA ET AL.
102
Massarinaceae Munk, Friesia 5: 305. 1956.
Type genus:Massarina Sacc.
Helminthosporium Link, Mag. Gesell. naturf. Freunde, Berlin
3(1–2): 10. 1809.
Type species: Helminthosporium velutinum Link.
Notes: Although more than 700 taxa have been described as
species within Helminthosporium (http://www.indexfungorum.org,
Aug. 2015), the genus has been quite heterogeneous. Several
unrelated pathogens of the Poaceae were segregated from
Helminthosporium to other genera, i.e., Bipolaris (= Cochliobo-
lus), Curvularia (= Pseudocochliobolus), Exserohilum (= Setos-
phaeria), and Pyrenophora (= Drechslera), all of which belong to
the Pleosporaceae (Sivanesan 1987, Hyde et al. 2013). Further
distantly related species (e.g., H. asterinum) in the Leotiomy-
cetes were excluded from Helminthosporium, and this genus
was restricted to species having phylogenetic affinity with the
Massarinaceae (Olivier et al. 2000, Kodsueb et al. 2007, Hyde
et al. 2013).
Little is known about the sexual morphs of Helminthospo-
rium s. str. Hughes (1953a) reported that an ascospore isolate
of a Massaria species found on Quercus produced a Hel-
minthosporium asexual morph. Subramanian & Sekar (1987)
described Splanchnonema kalakadense as the sexual morph
of H. velutinum based on cultural study. However, the validity
of the generic classification based on the sexual morphs
of these two examples remains unknown. The sexual morph
of H. massarinum differs from those of Massaria (Massar-
iaceae;Voglmayr & Jaklitsch 2011)andSplanchnonema
(Pleomassariaceae;Hyde et al. 2013) in its ellipsoidal hyaline
ascospores, and is rather similar to that of Massarina,
although M. eburnea (the type species of Massarina)does
not have a hyphomycetous asexual morph like that of
Helminthosporium.
Helminthosporium massarinum Kaz. Tanaka, K. Hiray. &
Shirouzu, sp. nov. MycoBank MB811315. Fig. 25.
Etymology: Referring to the similarity of the sexual morph with
that of the genus Massarina.
Ascomata numerous, scattered to 2–4 grouped, immersed
below the host epidermis, hemispherical to subglobose with
somewhat flattened base, 315–390 μm high, 300–430 μm diam.
Ostiolar neck central, cylindrical to papillate, 125–165 μm long,
75–125 μm wide, composed of subglobose, 3–5μm diam,
brown cells, surrounded by dark brown clypeus-like structure,
without periphyses. Ascomatal wall surface of textura prismatica,
in a longitudinal section 12–18 μm wide at sides and base,
composed of 5–7 layers of polygonal to rectangular,
5–15 × 2.5–6.5 μm, brown cells. Pseudoparaphyses cellular,
1.5–3μm wide, septate at 7–16 μm long intervals, branched,
anastomosed. Asci fissitunicate, clavate, 82–135 × 13–16 μm
(av. 111.0 × 14.8 μm, n = 90), rounded at the apex, with a narrow
apical chamber and faint ring, short-stalked (7–24 μm long; av.
16.7 μm, n = 35), with 8 ascospores biseriate above and uni-
seriate below. Ascospores ellipsoidal with rounded ends, mostly
straight, 20–25(–27) × 5–8μm (av. 22.6 × 7.0 μm, n = 100), l/w
2.7–3.9 (av. 3.2, n = 100), with a submedian septum (0.50–0.58;
av. 0.54, n = 100), constricted at the septum, asymmetric, with
wider upper cell, hyaline, guttulate, smooth, with a conspicuous
gelatinous sheath of 2–3μm wide in fresh condition (with a
delimited firm sheath of 1 μm wide in dry condition).
Colonies on PDA (after 4 wk) attaining a diam of 5.5–7.1 cm,
white to olivaceous grey; reverse smoke grey to buff; no
pigment produced. In culture hyphomycetous asexual morph
and spermatial morph formed. Conidiophores macronematous,
mononematous, 380–810 μm long (av. 587 μm, n = 20),
7–9μm wide at the apex, 13.5–21 μmwideatthebase,
arising singly or in groups of 4–5 from the stromata, straight
or slightly curved, dark brown, guttulate, 15–25-septate at
8–45 μm long intervals, with small (ca. 1–2μmdiam)poresat
the apex and beneath of septa, with thick wall of 1–4μmwide.
Conidia tretic, solitary or in short chains (5–6), 1–8-septate,
17–56.5 × 5–9μm (av. 37.9 × 7.4 μm, n = 95), l/w 2.9–7.5
(av. 5.1, n = 95) [but mostly 4–6-septate, 27 –53 × 6 –9μm
(av. 38.7 × 7.4 μm, n = 64), l/w 4.1–6.8(av.5.3,n=64)],
obclavate, rostrate, pale brown, smooth, with or without gut-
tules. Conidia germinating from both end cells. Spermogonia
pycnidial, produced under the conidiophores, solitary
or gregarious, 100–150( –300) μm diam, subglobose, uni-
loculate to multiloculate. Spermatiogenous cells cylindrical, up
to 5 μm long, appearing phialidic. Spermatia 3–4(–5) ×
2–2.5 μm(av.3.5×2.2μm, n = 26), globose to subglobose,
hyaline, smooth.
Specimens examined:Japan, Aomori, Towada, Sanbongi, Yagami, on vines of
Berchemia racemosa, 15 Dec. 2001, S. Hatakeyama, KT 838 = HHUF 27573,
culture JCM 13094 = MAFF 239604; ibid., 23 Nov. 2003, S. Hatakeyama
D
A
G
CEF
B
Fig. 24. Tingoldiago graminicola. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G.
Ascospore (arrowheads indicate gelatinous sheath; in India ink). A–D, F, G from KH 68; E from culture KT 891. Scale bars: A = 1 mm; B = 500 μm; C = 100 μm; D–G=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 103
(holotype KT 1564 = HHUF 29089, culture ex-type CBS 139690 = JCM
13095 = MAFF 239605); ibid., KT 1565 = HHUF 29090; ibid., KT 1566 = HHUF
29091; ibid., 2 Dec. 2003, K. Tanaka, S. Hatakeyama & N. Nakagawara, KT
1613–1615 = HHUF 29092–29094.
Note: This species is somewhat similar to H. hypselodelphyos in
having solitary or catenate conidia but the latter has smaller
conidia (15–28 × 6.5–8μm, av. 25 × 7.1 μm) (Ellis 1961).
Massarina Sacc., Syll. Fung. 2: 153. 1883.
Type species:Massarina eburnea (Tul. & C. Tul.) Sacc.
Notes: Since establishment of the genus (Saccardo 1883), more
than 176 taxa have been described within Massarina (Index
Fungorum; http://www.indexfungorum.org, Aug. 2015), but the
heterogeneity of the genus has been continuously suggested.
Several taxonomic reassessments of many species within
Massarina have been attempted based on morphology (Bose
1961, Barr 1992, Hyde 1995, Aptroot 1998, Hyde et al. 2002,
Tanaka & Harada 2003b). It has more recently been revealed
that the genus is highly polyphyletic based on molecular data
(Liew et al. 2002), and most species in Massarina except for the
type (M. eburnea) have been excluded from the genus. The
genera segregated from Massarina s. lat. are phylogenetically
diverse groups in the Pleosporales;Halomassarina (Trem-
atosphaeriaceae;Suetrong et al. 2009), Lentithecium and Tin-
goldiago (Lentitheciaceae;Zhang et al. 2009c, Hirayama et al.
2010), Lindgomyces (Lindgomycetaceae;Hirayama et al.
2010), Morosphaeria (Morosphaeriaceae;Suetrong et al.
2009), and Triplosphaeria (Tetraplosphaeriaceae;Tanaka et al.
2009).
Massarina eburnea (Tul. & C. Tul.) Sacc., Syll. Fung. 2: 153.
1883. Fig. 26.
A
N
D
O
CB
E
FG
I
JK
MPQ
RS
TU
L
H
Fig. 25. Helminthosporium massarinum. A, B. Ascomata on the natural host surface; C, D. Ascomata in longitudinal section (in lactophenol cotton blue); E. Ascomatal wall (in
lactophenol cotton blue); F. Pseudoparaphyses; G. Ascus; H–J. Ascospores (arrowheads indicate gelatinous sheath; J. in Black-Blue ink); K. Germinating ascospore; L–O.
Conidiophores; P. Conidiogenous cells (arrows indicate tretic pores); Q–S. Conidia; T. Spermogonium; U. Spermatia. A–I, K from KT 1564; J from KT 838; L–U from culture KT
838. Scale bars: A = 2 mm; B, L, M, T = 200 μm; C, D, N, O = 50 μm; E–K, P–S, U = 10 μm.
TANAKA ET AL.
104
Basionym:Massaria eburnea Tul. & C. Tul., Select. Fung. Car-
pol. 2: 239. 1863.
Ascomata scattered to 2–4 grouped, immersed, hemispherical
with flattened base or depressed globose, 300–420 μm high,
570–680 μm diam. Ostiolar neck short papillate, 60–90 μm long,
central, with black clypeus (250–400 μm wide). Ascomatal wall
12–18 μm thick at sides, composed of 3–5 layers of thin-walled
prismatic cells (5–12.5 × 2–4μm). Pseudoparaphyses
numerous, cellular, 2–3(–4.5) μm wide. Asci fissitunicate,
clavate, (110–)120–180 × 21.5–30 μm (av. 141.6 × 25.6 μm,
n = 50), with a stipe of 12.5–25 μm long (av. 18.6 μm, n = 38).
Ascospores broadly fusiform with rounded ends,
34–40 × 12–15(–16) μm (av. 37.2 × 13.9 μm, n = 50; including
firm sheath of 1–2μm thick), l/w 2.5–2.9 (av. 2.7, n = 50), 3-
septate (primary septum submedian: 0.51–0.55, av. 0.53,
n = 50), hyaline, smooth, with a sharply delimited firm sheath of
1–2μm thick (but up to 6 μm thick when fresh).
Specimen examined:UK, Wales, Swansea, dead twigs of Fagus sylvatica, Apr.
2001, H 3953 = HHUF 26621, culture CBS 139697 = JCM 14422.
Notes: The morphological characteristics of the above specimen
fit well with those in the description of M. eburnea (Hyde 1995). A
pycnidial morph of M. eburnea reported as Ceratophoma sp.
(Bose 1961, Sivanesan 1984) was observed in our isolate, but
the “conidia”did not germinate on several agar media (e.g.,
water agar, PDA, CMA, and MEA) over 7 d. The Ceratophoma
morph is therefore considered to be spermatial in function.
Stagonospora (Sacc.) Sacc., Syll. Fung. 3: 445. 1884.
Type species:Stagonospora paludosa (Sacc. & Speg.) Sacc.
Notes:Stagonospora has been defined morphologically based
on a broad generic concept (Sutton 1980) and has been believed
to have phylogenetic affinities with the Phaeosphaeriaceae
(Zhang et al. 2012). However, in a comprehensive phylogenetic
study on Stagonospora and morphologically similar genera,
Quaedvlieg et al. (2013) revealed that Stagonospora is poly-
phyletic and Stagonospora s. str. (based on S. paludosa) be-
longs to the Massarinaceae. Several new genera, such as
Neostagonospora and Parastagonospora, have been estab-
lished to accommodate unrelated stagonospora-like species in
the Phaeosphaeriaceae (Quaedvlieg et al. 2013).
Neottiosporina, typified by N. apoda (Subramanian 1961)
may have phylogenetic relationships with Stagonospora,
although phylogenetic placement of the type species has not
been clarified. At least N. paspali, transferred from Stagonospora
to Neottiosporina (Sutton & Alcorn 1974), should be regarded
within Stagonospora, because this taxon has close morpholog-
ical and phylogenetic affinity to Stagonospora species (Fig. 1).
Stagonospora bicolor (D. Hawksw. et al.) Kaz. Tanaka & K.
Hiray., comb. nov. MycoBank MB811316.
Basionym:Leptosphaeria bicolor D. Hawksw. et al., Mycologia
71: 483. 1979.
≡Saccharicola bicolor (D. Hawksw. et al.) D. Hawksw. & O.E. Erikss.,
Mycologia 95: 431. 2003.
Notes: Although we did not observe any material of this species,
the illustration in the original description of Leptosphaeria bicolor
(Kaiser et al. 1979), as well as SSU (GenBank U04202) and ITS
(GenBank U04203) sequences derived from the type culture
(ATCC 42652) of this species clearly indicates that it is a member
of Stagonospora, as recently circumscribed by Quaedvlieg et al.
(2013). Based on L. bicolor,Eriksson & Hawksworth (2003)
erected Saccharicola for leptosphaeria-like species on sugar-
cane, but Saccharicola should be regarded as a synonym of
Stagonospora. The sexual morph of Stagonospora s. str. has
been referred to as didymella-like (Quaedvlieg et al. 2013), and
some species of Didymella with a stagonospora-like asexual
morph (e.g., D. proximella on Carex;Corlett & Smith 1978) may
have phylogenetic affinity with Stagonospora s. str. based on
their morphologies.
Stagonospora perfecta Quaedvlieg et al., Stud. Mycol. 75: 378.
2013. Fig. 27.
D
K
AB
E
C
FJ
GL
HI
Fig. 26. Massarina eburnea. A, B. Ascomata on the natural host surface; C. Side view of ascoma; D. Ascoma in longitudinal section; E. Ascomatal wall; F. Pseudoparaphyses;
G. Ascus; H–J. Ascospores (arrowheads indicate gelatinous sheath); K. Spermogonia in culture; L. Spermatia. A –J from H 3953; K, L from culture H 3953. Scale bars:
A = 1 mm; B, C, K = 500 μm; D = 100 μm; E–J, L = 10 μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 105
Ascomata scattered, immersed, globose to subglobose in
section, 160–220 μm high, 140–250 μmdiam.Ostiolar neck
central, papillate to short cylindrical, 37–50 μm long,
52–75(–100) μmwide.Ascomatal wall in longitudinal section
uniformly 12–20 μm thick, composed of 4–6layersof
polygonal to subglobose, brown cells of 5–20 × 3.5–10 μm.
Pseudoparaphyses cellular, septate, branched and anasto-
mosed, 2.5–4μmwide.Asci fissitunicate, cylindrical,
75–115 × 15 –20.5 μm (av. 89.8 × 17.5 μm, n = 30), rounded
at the apex and with a shallow ocular chamber, short-stalked
(7–14.5 μm long), with 8 biseriate ascospores. Ascospores
broadly fusiform, 20–28 × 6.5–11 μm (av. 23.4 × 8.3 μm,
n=50),l/w2.3–3.3 (av. 2.8, n = 50), with a submedian
septum (0.54–0.59; av. 0.56, n = 50), constricted at
the septum, hyaline, smooth, with an entire sheath;
sheath gelatinous, 2–7μm, staining with Black-Blue ink when
in fresh condition, delimited and 1–3μmwidewhenindry
condition.
In culture both sexual and asexual morphs formed. Conidiomata
pycnidial, 80–100 μm high, 100–120 μm diam, scattered,
globose in section. Conidiomatal wall in longitudinal section
uniformly 6–13 μm wide, composed of 3–4 layers of polygonal,
thin-walled, 7–17 × 2.5–6μm, pale brown cells. Conidiophores
absent. Conidiogenous cells annellidic, cylindrical to subglobose,
8–17 × 5–9μm. Conidia cylindrical, 3(–5)-septate, hyaline,
smooth, (25–)27–37(–48) × 8–11 μm (av. 32.9 × 9.5 μm,
n = 60), l/w 2.7–4.1(–5.1) (av. 3.5, n = 60), without sheath.
Ascospores in culture slightly larger than those on the natural
host, 27–33 × 10–12 μm.
Specimen examined:Japan, Hokkaido, Isl. Rebun, Funadomari, Kusyuko (pond),
on dead leaves of Carex sp., 3 Jun. 2004, K. Tanaka, KT 1726A = HHUF 29095,
culture JCM 13099 = MAFF 239609.
Notes: The characteristics of the above material match those in
the original description of S. perfecta described from Carex
acutiformis in the Netherlands (Quaedvlieg et al. 2013), except
that the conidia in our material were wider (8–11 μmvs.
6–8μm). LSU and ITS sequences obtained from our material
and those from the ex-type of S. perfecta (GenBank LSU:
KF251761, ITS: KF251258) were identical. The sexual morph of
this species has been referred to as didymella-like (Quaedvlieg
et al. 2013), and fits well with the generic concept of Saccharicola
(Eriksson & Hawksworth 2003), but this genus should be syn-
onymised under the older genus Stagonospora (Saccardo 1884).
Stagonospora pseudoperfecta Kaz. Tanaka & K. Hiray., sp.
nov. MycoBank MB811317. Fig. 28.
Etymology: After its morphological similarity to Stagonospora
perfecta.
Ascomata scattered, immersed, globose in section, 200–250 μm
high, 210–260 μm diam. Ostiolar neck central, papillate,
35–63 μm long, 63 μm wide. Ascomatal wall in longitudinal
section uniformly 10–15 μm thick, composed of 3–5 layers of
polygonal to subglobose, brown cells of 6–16.5 × 3.5–11.5 μm.
Pseudoparaphyses cellular, numerous, 3–6μm wide, septate,
branched, anastomosed. Asci fissitunicate, clavate,
66–100 × 13.5–17.5 μm (av. 86.3 × 15.4 μm, n = 20), rounded at
apex and with a shallow apical chamber, short-stalked
(7–17.5 μm long), with 8 irregularly biseriate to triseriate asco-
spores. Ascospores narrowly fusiform, straight or slightly curved,
21–30.5 × 5–7μm (av. 26.0 × 5.7 μm, n = 40), l/w 3.3–5.9 (av.
4.6, n = 40), with submedian septum (0.50–0.57, av. 5.4, n = 40),
slightly constricted at the septum, hyaline, with or without gut-
tules, smooth, with an entire sheath; sheath gelatinous,
0.5–2μm wide at side.
I
E
FH
BA
G
JK
C
D
NO
Q
P
M
L
Fig. 27. Stagonospora perfecta. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall in surface view; E. Ascomatal wall at side; F.
Pseudoparaphyses; G. Asci; H. Apex of ascus; I–K. Ascospores (arrowheads indicate gelatinous sheath; I. in Black-Blue ink; K. in India ink); L, M. Conidiomata in culture (on
rice straw); N, O. Conidiogenous cells (arrowheads indicate annellations); P. Conidioma in longitudinal section; Q. Conidia. A–K from KT 1726; L–Q from culture KT 1726.
Scale bars: A, L = 1 mm; B, M = 200 μm; C, P = 20 μm; D –K, N, O, Q = 10 μm.
TANAKA ET AL.
106
Colonies on PDA (after 2 wk) attaining a diam of 4.9–5.5 cm,
white; reverse sepia to fuscous black; no pigment produced. In
culture asexual morph formed. Conidiomata pycnidial. Conidia
cylindrical, aseptate, hyaline, 21.5–26 × 4–5.5 μm (av.
24.1 × 4.9 μm, n = 10), l/w 4.6–5.8 (av. 5.0, n = 10).
Specimens examined:Japan, Aomori, Hirosaki, Kadoke, Oowasawa-river,
riverbank (Horikoshi-bridge), on dead leaves of Typha latifolia, 8 Sep. 2002, K.
Tanaka (holotype KT 889 = HHUF 29087, culture ex-type CBS 120236 = JCM
13097 = MAFF 239607); ibid., KT 888 = HHUF 29086.
Notes: Eight species are currently accepted in Stagonospora s.
str. based on molecular evidence (Crous et al. 2013b, 2014b,
Quaedvlieg et al. 2013). Stagonospora pseudoperfecta is
similar to S. perfecta, but has slightly longer and more slender
ascospores (l/w 4.6 vs. 2.8). In terms of overall morphology and
host preference, S. pseudoperfecta superficially resembles
“Massarina”lacustris sensu Leuchtmann (1984) (non Wett-
steinina lacustris sensu Shoemaker & Babcock 1989,or
Khashnobish & Shearer 1993) reported from Typha and
Schoenoplectus. However, molecular data (SSU, rpb2,tef1)
obtained from a strain studied by Leuchtmann (CBS 618.86)
suggests that the latter fungus is a member of the Lentithe-
ciaceae (Schoch et al. 2009). In ITS analysis with other Sta-
gonospora species (data not shown), S. pseudoperfecta
positioned as a sister taxon to S. duoseptata, but similarity
between the two taxa in this region was 95.2 % (452/475) with
1.3 % (6/475) gaps.
In culture, pycnidial conidiomata with cylindrical hyaline
conidia were observed only once, but this may have been an
immature condition because the conidia were aseptate and
smaller (Fig. 28J). Unfortunately, the asexual morph in culture
failed to be observed again, despite several attempts.
Stagonospora tainanensis W.H. Hsieh, Mycologia 71: 893.
1979. Fig. 29.
Asci ovoid to cylindrical, 102–122.5 × 26.5–32.5 μm, 8-
spored. Ascospores fusiform, 36 –44 × 8.5–12 μm, l/w
3.6–4.7, with a submedian septum (0.53–0.56), hyaline,
smooth,withanentiresheath(2–5μm wide when fresh,
1–2μmwidewhendry).
In culture, both sexual and asexual morphs formed. Ascospores
32–40 × 10.5–13 μm, l/w 2.8–3.2, with a submedian septum
(0.54–0.56). Conidia ellipsoid, 37–48(–55) × 12–13(–15) μm, l/
w 3.0–3.8, 3-septate, hyaline.
Specimen examined:Japan, Kagoshima, Isl. Yakushima, Nunobikinotaki park,
on dead leaves of herbaceous plant, 19 Oct. 2005, K. Tanaka & T. Hosoya, KT
1866 = HHUF 30141, culture MAFF 243860.
Notes: The ITS sequence from this material is identical with
the sequence (GenBank AF439464) of Stagonospora taiwa-
nensis obtained from the ex-type culture (ATCC 38204; Hsieh
1979), and two ITS sequences (GenBank AF439462,
AF439463) of Saccharicola taiwanensis on Saccharum.An
unnamed “Saccharicola”on Miscanthus (O'Neill & Farr 1996),
a grass genus related to Saccharum,isalsoconsideredtobe
conspecific, because these have identical ITS sequences
(GenBank AF439467; C^
amara et al. 2002) and morphological
similarities (O'Neill & Farr 1996). Morphologically, our
specimen agrees with the previous description of this species
(as Leptosphaeria taiwanensis;Hsieh 1979, Sivanesan
1984, Sivanesan & Waller 1986), but the large
ascospores reported by Shoemaker & Babcock (1989) were
not observed.
Morosphaeriaceae Suetrong et al., Stud. Mycol. 64: 161. 2009.
Type genus:Morosphaeria Suetrong et al.
Aquilomyces D.G. Knapp et al., Persoonia 35: 93. 2015.
Type species:Aquilomyces patris D.G. Knapp et al.
Notes:Aquilomyces was erected by Knapp et al. (2015) to
accommodate A. patris, a root endophyte of white poplar. There
was no morphological information of this genus, because no
fructifications were observed for A. patris.
Aquilomyces rebunensis Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811322. Fig. 30.
Etymology: After the locality where this fungus was collected.
C
AB
JI
DE
G
F
H
Fig. 28. Stagonospora pseudoperfecta. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus;
G–I. Ascospores (arrowheads indicate gelatinous sheath); J. Conidia. A–I from KT 889; J from culture KT 889. Scale bars: A = 2 mm; B = 500 μm; C = 20 μm; D –J=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 107
Ascomata subglobose, 350–550 μm high, (430–)600–700 μm
diam, covered with numerous brown hyphae of 2–3μm thick.
Ostiolar neck terete, central, 85–130 μm long, 75–130 μm wide,
clypeate, with periphyses, composed of carbonaceous polygonal
cells of 5–10 × 3–5μm. Ascomatal wall at side wedge-shaped,
up to 125 μm thick, composed of vertically orientated angular
brown cells (5–8×2–5μm); wall at the base 25–50 μm thick,
composed of 3–8μm diam, subglobose, brown cells. Pseudo-
paraphyses branched and anastomosed, septate, 1.5–2μm
wide. Asci fissitunicate, cylindrical to clavate, 97.5–147.5 ×
15–21 μm (av. 121.3 × 18.3 μm, n = 45), with a short stipe of
7.5–25(–37.5) μm (av. 18.0 μm, n = 34). Ascospores fusiform,
slightly curved, 30–38.5 × 6.5–11.5 μm (av. 35.1 × 8.6 μm,
n = 50), l/w 3.3–5.0(–5.6) (av. 4.2, n = 50), with a supramedian
septum (0.44–0.50; av. 0.47, n = 57), hyaline, smooth, with a
sheath of 1–2μm wide.
Colonies on PDA (after 4 wk) attaining a diam of 4.3–4.6 cm,
pale olivaceous grey with white margin; reverse black to
olivaceous grey; no pigment produced. In culture spermatial and
ascomatal morphs formed. Spermatia 2–3 × 1.5–1.8 μm, sub-
globose to oblong. Ascospores slightly larger than those found
on nature, 35–48 × 8–11 μm (av. 39.7 × 9.3 μm, n = 25).
Specimen examined:Japan, Hokkaido, Isl. Rebun, Kafuka, Nairo-river, on
submerged twigs of woody plant, 30 Aug. 2001, K. Tanaka (holotype KT 732-
2 = HHUF 27556, culture ex-type CBS 139684 = JCM 19427 = MAFF 243862).
Notes: We tentatively describe this aquatic fungus as Aquilo-
myces based on our molecular results. Phylogenetic analysis
showed that A. patris, the type species of Aquilomyces, and
A. rebunensis clustered into a distinct and moderately-supported
clade (85 %, Fig. 1). Sequence similarities between these two
taxa were 97.7 % (858/878) with 2.3 % gaps (20/878) in LSU and
95.4 % (576/604) with 0.7 % gaps (4/604) in ITS, suggesting that
they are congeneric.
Clypeoloculus Kaz. Tanaka & K. Hiray., gen. nov. MycoBank
MB811318.
I
G
FE
B
HD
CLKJ
A
Fig. 30. Aquilomyces rebunensis. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section (in lactophenol cotton blue); D. Ascomatal wall; E. Pseu-
doparaphyses; F. Ascus; G–K. Ascospores (arrowheads indicate gelatinous sheath); L. Spermatia. A–D, F–I from KT 732; E, J–L from culture KT 732. Scale bars: A = 1 mm;
B = 500 μm; C = 100 μm; D–L=10μm.
FD
ABC
K
EGJ
HI
Fig. 29. Stagonospora tainanensis. A, B. Ascomata on the natural host surface; C. Ascomata and conidiomata in culture (on rice straw); D. Ascoma in longitudinal section; E.
Ascomatal wall; F. Pseudoparaphyses; G. Ascus; H–J. Ascospores (arrowheads indicate gelatinous sheath); K. Conidia. A, B, D, E, G, H from KT 1866; C, F, I–K from culture
KT 1866. Scale bars: A, C = 2 mm; B = 500 μm; D = 20 μm; E–K=10μm.
TANAKA ET AL.
108
Etymology: Referring to the morphology of the ascomata covered
by clypeus.
Ascomata scattered to 2–3 grouped, immersed to erumpent,
subglobose or hemispherical with flattened base in section,
covered with numerous brown hyphae around ascomata.
Ostiolar neck terete to papillate, central, clypeate, composed of
carbonaceous cells. Ascomatal wall wedge-shaped or “rim-like”
at sides. Pseudoparaphyses septate, branched and anasto-
mosed. Asci fissitunicate, clavate, with a short stipe, 8-spored.
Ascospores narrowly fusiform, slightly curved, with a primary
septum median to supramedian, constricted at the septum, hy-
aline, smooth, surrounded by an entire gelatinous sheath.
Asexual morph unknown.
Type species:Clypeoloculus akitaensis Kaz. Tanaka & K. Hiray.
Notes: A new genus, Clypeoloculus, is proposed here for fresh-
water species having globose to subglobose ascomata with
prominent clypeus, clavate asci, and 1-septate, hyaline asco-
spores with an entire sheath. These morphological characters fit
with the broad generic concept of Massarina (Aptroot 1998,
Tanaka & Harada 2003b), but Clypeoloculus can be distin-
guished from Massarina s. str. by the ascomata with wedge-
shaped or “rim-like”ascomatal wall and narrowly fusiform asco-
spores. Furthermore, species in Clypeoloculus are located in the
Morosphaeriaceae clade and are distantly related to M. eburnea,
the type species of Massarina (Massarinaceae;Fig. 1).
Clypeoloculus akitaensis Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811319. Fig. 31.
Etymology: Referring to the location where the specimen was
collected.
Ascomata scattered to 2–3 grouped, immersed to erumpent,
globose to subglobose in section, 400–550 μm high,
580–720 μm diam, covered with numerous brown hyphae
around ascomata. Ostiolar neck terete, central, clypeate,
composed of carbonaceous cells. Ascomatal wall in longitudinal
section 37–100 μm wide at sides and base, composed of sub-
globose to globose, thick-walled, 2.5–10 μm diam, brown cells.
Pseudoparaphyses septate, branched and anastomosed,
2–2.5 μm wide. Asci fissitunicate, clavate, (95–)
110 –155 × 17.5–27.5 μm (av. 125.1 × 20.9 μm, n = 30 μm), with
a short stipe of (7.5–)12–27.5(–30) μm (av. 19.4 μm, n = 26)
long, with 8 biseriate ascospores. Ascospores narrowly fusiform,
slightly curved, 33.5–43 × 7–10 μm (av. 38.7 × 8.6 μm, n = 50), l/
w 3.8–5.2 (av. 4.5, n = 50), with a primary septum supramedian
(0.44–0.50; av. 0.48, n = 47), constricted at the septum, hyaline,
smooth, surrounded by an entire gelatinous sheath, 1.5–3μm
wide.
Colonies on PDA (after 4 wk) attaining a diam of 2.0–2.2 cm,
greenish grey; reverse black to smoke grey; no pigment pro-
duced. Spermatial morph observed in culture. Spermatia oblong,
hyaline, 2.8–4 × 1.3–1.9 μm (av. 3.3 × 1.7 μm, n = 30).
Specimen examined:Japan, Akita, Kisakata, Akagawa-river, riverbank, on
submerged twigs of woody plant, 23 Sep. 2001, K. Tanaka (holotype KT
788 = HHUF 27557, culture ex-type CBS 139681 = JCM 19424 = MAFF
239467).
Notes: In ascospore dimensions, C. akitaensis (av.
38.7 × 8.6 μm) is most similar to C. towadaensis (av.
38.7 × 8.7 μm), but the latter species has hemispherical asco-
mata with flattened bases and longer asci [(112–)120 –170 μm
long]. In the ITS sequences of these two species, there were
differences at 29 positions.
Tanaka & Harada (2003b) reported this fungus as Massarina
peerallyi (Hyde & Aptroot 1998). However, our re-examination of
M. peerallyi based on its holotype [HKU (M) 2409] revealed that
they are distinct species, because M. peerallyi has broader asci
(80–127.5 × 25–35 μm) and larger ascospores
(37–45 × 9 –12 μm) with a supramedian septum (0.43–0.50 av.
0.48, n = 32). The morphological features of M. peerallyi are
generally in accordance with the generic concept of Clypeoloculus,
but phylogenetic reassessment using molecular sequences will be
necessary before a new combination is proposed for this species.
A
F
B
ELK
GHI
D
C
J
Fig. 31. Clypeoloculus akitaensis. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G–I.
Ascospores (arrowheads indicate gelatinous sheath); J. Spermogonium in culture (on rice straw); K. Spermogonium in longitudinal section (in lactophenol cotton blue); L.
Spermatia. A–I from KT 788; J–L from culture KT 788. Scale bars: A = 2 μm; B, J = 500 μm; C, K = 100 μm; D –I, L = 10 μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 109
Clypeoloculus hirosakiensis Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811320. Fig. 32.
Etymology: Referring to the collection site.
Ascomata scattered, immersed to erumpent, globose to sub-
globose in section, 350–520 μm high, 340–550 μm diam,
covered with numerous brown hyphae around ascomata.
Ostiolar neck terete, central, 75–100 μm long, 40–170 μm
wide, clypeate, composed of carbonaceous cells. Ascomatal
wall in longitudinal section at sides, 50–60 μm wide, composed
of parallel rows of rectangular to polygonal, brown to dark
brown cells of 7.5–12.5 × 3.5–9.5 μm; wall at the base
17–25 μm wide, composed of subglobose to polygonal, slightly
thick-walled, 3–10 × 3.5–6.5 μm, brown cells. Pseudopar-
aphyses septate, branched and anastomosed, 1.5–3μm wide.
Asci fissitunicate, clavate, 100–147.5 × (13.5–)15–20 μm (av.
118.6 × 16.5 μm, n = 50 μm), with a short stipe of (7.5–)
10–25(–30) μm (av. 16.5 μm, n = 50) long, with 8 biseriate
ascospores. Ascospores narrowly fusiform, slightly curved,
(24–)29–36 × 7–9.5 μm (av. 33.0 × 8.0 μm, n = 50), l/w
3.7–4.7 (av. 4.1, n = 50), with a primary septum nearly median
(0.47–0.52; av. 0.50, n = 50), constricted at the septum, hya-
line, smooth, surrounded by an entire gelatinous sheath,
3–7μm wide.
Colonies on PDA (after 4 wk) attaining a diam of 2.7–3.0 cm,
greenish grey to lavender grey; reverse chestnut; ochreous
pigment produced. In culture spermatial morph formed. Sper-
matia oblong, hyaline, 2.6–3.5 × 1.2–1.5.
Specimen examined:Japan, Aomori, Hirosaki, Zatoishi (river), on submerged
twigs of woody plant, 19 Jul. 2003, K. Tanaka & N. Asama (holotype KT
1283 = HHUF 30144, culture ex-type CBS 139682 = JCM 19425 = MAFF
243864).
Note: The ascospores of C. hirosakiensis (av. 33.0 × 8.0 μm) are
larger than those of C. microsporus (av. 29.9 × 7.0 μm) but
smaller than those of C. akitaensis (av. 38.7 × 8.6 μm) and
C. towadaensis (av. 38.7 × 8.7 μm).
Clypeoloculus microsporus Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811321. Fig. 33.
Etymology: Referring to the small ascospores.
Ascomata scattered to 2–3 grouped, immersed to erumpent,
globose to subglobose in section, 290–310 μm high,
280–350 μm diam, covered with sparse brown hyphae around
ascomata. Ostiolar neck central, papillate, 70–90 μm long,
100–120 μm wide, clypeate, composed of carbonaceous cells.
Ascomatal wall in longitudinal section 25–38 μm wide at sides
and 25–30 μm wide at the base, composed of subglobose brown
cells (2.5–10 μm diam). Pseudoparaphyses branched and
anastomosed, septate, 2–3μm wide. Asci fissitunicate, cylin-
drical to clavate, 80–145 × 13.5–22.5 μm (av. 115.5 × 17.5 μm,
n = 33), with a short stipe of (5–)7.5–15(–29.5) μm (av. 13.9 μm,
n = 30) long, with 8 biseriate to triseriate ascospores. Asco-
spores narrowly fusiform, slightly curved,
25.5–34.5 × 6–8(–9) μm (av. 29.9 × 7.0 μm, n = 56), l/w
3.4–5.2(–5.5) (av. 4.3, n = 56), with a primary septum nearly
median (0.47–0.52; av. 0.50, n = 56), constricted at the septum,
hyaline, smooth, surrounded by an entire gelatinous sheath of
2–4μm wide.
Colonies on PDA (after 4 wk) attaining a diam of 3.1–3.3 cm,
olivaceous grey; reverse umber to chestnut; amber pigment
produced. In culture spermatial morph formed. Spermatia
oblong, hyaline, 2.5–3.2 × 1.2–1.6 μm.
Specimen examined:Japan, Aomori, Kuroishi, Lake Nizinoko, on submerged
twigs of woody plant, 3 May 2003, K. Tanaka & N. Asama (holotype KT
1131 = HHUF 30143, culture ex-type CBS 139683 = JCM 19426 = MAFF
243863).
Note: This species is distinguished from other species of Cly-
peoloculus by having the smallest ascospores known in the
genus.
Clypeoloculus towadaensis Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811323. Fig. 34.
B
E
D
FJKL
C
I
A
H
G
Fig. 32. Clypeoloculus hirosakiensis. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G–I.
Ascospores (arrowheads indicate gelatinous sheath; I. in India ink); J. Germinating ascospore; K. Spermogonia in culture (on rice straw); L. Spermatia. A–J from KT 1283; K, L
from culture KT 1283. Scale bars: A = 1 mm; B, K = 500 μm; C = 100 μm; D–J, L = 10 μm.
TANAKA ET AL.
110
Etymology: Referring to the collection site.
Ascomata scattered to 2–3 grouped, immersed to erumpent,
hemispherical with flattened base in section, 260–360 μm high,
550–700 μm diam. Ostiolar neck central, terete, 65–135 μm
long, 75–88 μm wide, clypeate, composed of carbonaceous
cells. Ascomatal wall at sides 95–150 μm wide and “rim-like”,
composed of vertically orientated, rectangular to subglobose,
hyaline to pale brown cells of 6–10 × 3–6μm; at the base
15–25 μm wide or poorly developed. Pseudoparaphyses
septate, branched and anastomosed, 1–2μm wide. Asci fissi-
tunicate, cylindrical, (112–)120–170 × 17.5 –25 μm (av.
136.7 × 21.3 μm, n = 35), with a short stipe of (7.5–)12.5–25 μm
(av. 17.2 μm, n = 35) long, with 8 biseriate ascospores. Asco-
spores narrowly fusiform, slightly curved, 34–43 × 7–10 μm (av.
38.7 × 8.7 μm, n = 50), l/w 3.9–5.3 (av. 4.5, n = 50), with a
primary septum nearly median (0.47–0.53; av. 0.50, n = 50),
constricted at the septum, hyaline, smooth, surrounded by an
entire gelatinous sheath of 1–5μm wide. Senescent ascospores
becoming brown, 1–3(–5)-septate.
Colonies on PDA (after 4 wk) attaining a diam of 1.9–2.2 cm,
greenish grey; reverse buff to sepia; ochreous pigment pro-
duced. In culture spermatial morph formed. Spermatia oblong to
subglobose, hyaline, 3–4 × 1.4–1.8 μm.
Specimen examined:Japan, Akita, Kazuno, Kosaka, Lake Towada, on sub-
merged twigs of woody plant, 26 Jul. 2003, K. Tanaka &N. Asama (holotype KT
1340 = HHUF 30145, culture ex-type CBS 139685 = JCM 19428 = MAFF
243865).
Notes: The most distinctive feature of C. towadaensis is its
hemispherical ascomata with “rim-like”ascomatal walls. The asci
B
EFLJ
GH
D
A
K
I
C
Fig. 33. Clypeoloculus microsporus. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G –J.
Ascospores (arrowheads indicate gelatinous sheath); K. Spermogonium in culture (on rice straw); L. Spermatia. A–J from KT 1131; K, L from culture KT 1131. Scale bars:
A = 1 mm; B, K = 500 μm; C = 100 μm; D–J, L = 10 μm.
A
EFLKJ
B
HI
D
C
G
Fig. 34. Clypeoloculus towadaensis. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G–J.
Ascospores (arrowheads indicate gelatinous sheath; J. senescent ascospore); K. Spermogonia in culture; L. Spermatia. A–J from KT 1340; K, L from culture KT 1340. Scale
bars: A = 1 mm; B, K = 500 μm; C = 100 μm; D–J, L = 10 μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 111
of this species are longer than those of other species in
Clypeoloculus.
Helicascus Kohlm., Canad. J. Bot. 47: 1471. 1969.
Type species:Helicascus kanaloanus Kohlm.
Notes:Helicascus was revised by Zhang et al. (2013) who
accepted six species: two marine (H. kanaloanus and
H. nypae) and four freshwater species (H. aegyptiacus,
H. aquaticus,H. elaterascus and H. thalassioideus). Recently,
two additional species in freshwater habitat (H. gallicus and
H. unilocularis) were described in the genus (Zhang et al.
2014b, 2015).
Helicascus aquaticus H. Zhang & K.D. Hyde, Sydowia 65: 155.
2013. Fig. 35.
Ascomata scattered to 2–3 grouped, immersed, hemispherical in
section, 170–300 μm high (excluding the long ostiolar neck),
230–530 μmdiam.Ostiolar neck cylindrical, oblique to central,
230–340 μmlong,110–160 μm wide, composed of polygonal to
subglobose, thick-walled, 2.5–7.5 × 2–4μm, brown to dark brown
cells, with dense periphyses. Ascomatal wall in longitudinal section,
40–110 μmthickatsides,“rim-like”, composed of parallel rows of
rectangular to polygonal, brown cells of 3.5–20 × 2.5–10 μm; at
base poorly developed, 7–15 μm thick, composed of thin-walled
flattened cells. Pseudoparaphyses septate, branched and anas-
tomosed, 1.5–2(–3.5) μm wide, associated with gelatinous mate-
rial. Asci fissitunicate, cylindrical to clavate, 80–122.5 ×
15–19.5 μm(av.102.8×17.5μm, n = 20), with a stipe of
17.5–32.5 μm long (av. 24.7 μm long, n = 20), 8-spored. Asco-
spores broadly fusiform, slightly curved, 21.5–29 × 8 –10.5 μm(av.
24.4 × 9.1 μm, n = 50), l/w 2.2–3.3 (av. 2.7, n = 50), with a sub-
median septum (0.51 –0.60; av. 0.57, n = 50), constricted at the
septum, pale greenish brown to brown, smooth, without sheath.
In culture spermatial morph formed. Spermatia oblong, hyaline,
2.4–4 × 1.5–2μm.
Specimen examined:Japan, Aomori, Hirosaki, Aoki, Mohei-pond, on submerged
twigs of woody plant, 25 Oct. 2003, K. Tanaka & N. Asama, KT 1544 = HHUF
30146, culture JCM 19423 = MAFF 243866.
Notes: The ITS sequence from the above isolate showed 100 %
identity with that from the ex-type of H. aquaticus (GenBank
KC886639), an aquatic species formerly known only from the
type specimen collected on submerged wood in Thailand (Zhang
et al. 2013). Our collection has a longer ascomatal neck (vs.
130–200 μm) and shorter asci [vs.90–140( –185) μm] than
described in the original publication. The ascospores possessing
three septa were noted as the most distinctive feature of
H. aquaticus (Zhang et al. 2013), but these were consistently 1-
septate in our material.
Helicascus elaterascus (Shearer) H. Zhang & K.D. Hyde,
Sydowia 65: 158. 2013. Fig. 36.
Basionym:Kirschsteiniothelia elaterascus Shearer, Mycologia
85: 963. 1994 (1993).
Ascomata scattered, immersed, depressed globose in section,
250–300 μm high (excluding the long ostiolar neck),
HI
D
M
LJK
C
AB
E
O
N
FG
Fig. 35. Helicascus aquaticus. A, B. Ascomata on the natural host surface; C–E. Ascomata in longitudinal section; F. Ostiolar neck of ascoma; G. Ascomatal wall; H.
Pseudoparaphyses; I. Ascus; J–L. Ascospores; M. Germinating ascospore; N. Spermogonium in culture (on rice straw); O. Spermatia. A–M from KT 1544; N, O from culture
KT 1544. Scale bars: A = 1 mm; B, N = 500 μm; C–E = 100 μm; F, M = 20 μm; G–L, O = 10 μm.
TANAKA ET AL.
112
600–910 μm diam. Ostiolar neck 200–230 μm long,
110 –160 μm wide, with dense periphyses. Pseudoparaphyses
branched and anastomosed, septate, associated with gelatinous
material. Asci fissitunicate, clavate, ca. 140 × 19 μm, with a long
stipe, 8-spored. Ascospores broadly fusiform, slightly curved,
26.5–31.5 × 9–12.5 μm (av. 29.0 × 10.6 μm, n = 30), l/w 2.4–3.1
(av. 2.8, n = 30), with a submedian septum (0.50–0.58; av. 0.54,
n = 30), slightly constricted at the septum, pale greenish brown to
brown, with an entire sheath slightly enlarged below. In culture
only sexual morph observed.
Specimens examined:Japan, Okinawa, Isl. Ishigaki, Mt. Omoto (river), on
submerged twigs of woody plant, 2 Dec. 2009, Y. Kurihara, KT 2673 = HHUF
30147, culture MAFF 243867; Okinawa, Isl. Iriomote, Aira-river, on submerged
twigs of woody plant, 8, Dec. 2009, Y. Kurihara & A. Ooba, KT 2682 = HHUF
30451, culture CBS 139689.
Notes: The morphological features of the examined material
matched well with those in the original description of this species
(Shearer 1993,asKirschsteiniothelia elaterascus) and the LSU
sequences were identical with that of the species deposited in
GenBank (GenBank AY787934; Kodsueb et al. 2006). This
species has been reported frequently from submerged wood in
temperate and tropical freshwater habitats (Shearer & Raja
2010, Raja et al. 2011) including Japan (Tsui et al. 2003).
Helicascus thalassioideus (K.D. Hyde & Aptroot) H. Zhang &
K.D. Hyde, Sydowia 65: 159. 2013. Fig. 37.
Basionym:Massarina thalassioidea K.D. Hyde & Aptroot, Nova
Hedwigia: 498. 1998.
Asci clavate, 80–120 × 15–17.5 μm (av. 100.2 × 16.7 μm, n = 30),
with a stipe of 15–32.5 μm long (av. 23.0 μm, n = 30). Ascospores
25–31 × 8–10 μm (av. 27.5 × 8.6 μm, n = 30), l/w 2.8–3.5 (av. 3.2,
n = 30), with a median septum. Asexual morph unknown.
Specimens examined:Japan, Okinawa, Isl. Iriomote, Geta-river, on submerged
twigs of woody plant, 22 Nov. 2008, K. Hirayama & K. Tanaka, KH 242 = HHUF
30069, culture JCM 17526 = NBRC 107811. China, Hong Kong, Lam Tsuen river,
N.T. on submerged wood, culture CBS 110441 = JCM 14147.
Notes: A more detailed description of this species was provided
by Hirayama & Tanaka (2011a). This species was originally
described as Massarina (Hyde & Aptroot 1998) and has
commonly been reported from submerged wood in freshwater
habitats throughout the world including Australia, Brunei, China
D
G
AB
M
J
I
C
H
F
L
K
E
Fig. 36. Helicascus elaterascus. A. Ascomata on the natural host surface; B. Ascomata in culture (on rice straw); C–E. Ascomata in longitudinal section; F. Ostiolar neck of
ascoma; G. Ascomatal wall; H. Pseudoparaphyses; I. Ascus; J–L. Ascospores (arrowheads indicate gelatinous sheath); M. Germinating ascospore. A, C–G, I, M from KT 2673;
B, H, J–L from culture KT 2673. Scale bars: A, B = 500 μm; C –E = 100 μm; F, M = 20 μm; G–L=10μm.
GFE
A
C
D
B
Fig. 37. Helicascus thalassioideus. A, B. Ascomata on the natural host surface; C. Ascomata in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G.
Ascospore. All from KH 242. Scale bars: A = 1 mm; B = 500 μm; C = 100 μm; D –G=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 113
(Yunnan), French West Indies, Hong Kong, the Philippines, and
Thailand (Zhang et al. 2013, 2015). Zhang et al. (2014b)
suspected that helicascus-like taxa with a worldwide distribu-
tion might be a species-complex. However, ITS sequences
obtained from the above two isolates were identical and differed
by only one position (517/518) from that of H. thalassioideus
collected in French West Indies (GenBank KP637162, Zhang
et al. 2015).
Morosphaeria Suetrong et al., Stud. Mycol. 64: 161. 2009.
Type species:Morosphaeria velatispora (K.D. Hyde & Borse)
Suetrong et al.
Notes: The genus Morosphaeria currently comprises two marine
species, M. ramunculicola and M. velatispora (Suetrong et al.
2009). A freshwater fungus, Kirschsteiniothelia elaterascus,
was transferred to Morosphaeria (Boonmee et al. 2012), but it is
presently placed in Helicascus (Zhang et al. 2013).
Morosphaeria ramunculicola (K.D. Hyde) Suetrong et al., Stud.
Mycol. 64: 162. 2009. Fig. 38.
Basionym:Massarina ramunculicola K.D. Hyde, Mycologia 83:
839. 1992 (1991).
Asci cylindrical to clavate, 121–187.5 × 22–32.5 μm, stalked
(20–42.5 μm long). Ascospores 33–41 × 11–15 μm (av.
36.5 × 12.6 μm, n = 30), l/w 2.4–3.4 (av. 2.9, n = 30), with a
submedian primary septum (0.50–0.56; av. 0.52, n = 30).
Asexual morph unknown.
Specimen examined:Japan, Okinawa, Isl. Iriomote, Oomijya-river, on twigs of
Rhizophora mucronata, 22 Nov. 2008, K. Hirayama & K. Tanaka, KH 220 = HHUF
30070, culture NBRC 107813.
Notes: The morphological features of the specimen cited here
were identical to those of M. ramunculicola (Hyde 1991).
Therewereonly1–2 bp differences between the LSU se-
quences (ca. 1 280 bp) of our material and those of
M. ramunculicola deposited in GenBank (GenBank
GQ925853, GQ925854).
Morosphaeria velatispora (K.D. Hyde & Borse) Suetrong et al.,
Stud. Mycol. 64: 161. 2009. Fig. 39.
Basionym:Massarina velatispora K.D. Hyde & Borse, Mycotaxon
27: 161. 1986.
Asci clavate, (135–)165–245(–280) × 25–38 μm (av.
212.2 × 32.8 μm, n = 30), with a stipe of 25–55 μm long (av.
37.0 μm, n = 26). Ascospores 42–52 × 14–18 μm (av.
47.1 × 16.1 μm, n = 40), l/w 2.6–3.4 (av. 2.9, n = 40), 1–3-
septate, with a median primary septum (0.49–0.53; av. 0.50,
n = 40). Asexual morph unknown.
Specimens examined:Japan, Okinawa, Isl. Iriomote, Oomijya-river, on twigs of
Rhizophora mucronata, 22 Nov. 2008, K. Hirayama & K. Tanaka, KH 218 = HHUF
30072, culture JCM 17529 = NBRC 107814; Okinawa, Isl. Iriomote, Geda river,
on twigs of Rhizophora mucronata, 20 Nov. 2008, K. Hirayama & K. Tanaka, KH
221 = HHUF 30073, culture JCM 17530 = NBRC 107812.
Note:Hirayama & Tanaka (2011a) provided a more precise
description based on the above specimens.
EGF
C
D
A
B
Fig. 38. Morosphaeria ramunculicola. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G.
Ascospore (arrowheads indicate gelatinous sheath). All from KH 220. Scale bars: A = 1 mm; B = 500 μm; C = 100 μm; D–G=10μm.
CEFG
D
B
A
Fig. 39. Morosphaeria velatispora. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G.
Ascospore (arrowheads indicate gelatinous sheath). A, D from KH 218; B, C, E–G from KH 221. Scale bars: A = 1 mm; B = 500 μm; C = 100 μm; D –G=10μm.
TANAKA ET AL.
114
Parabambusicolaceae Kaz. Tanaka & K. Hiray., fam. nov.
MycoBank MB811324.
Etymology: Referring to the name of the type genus.
Ascomata scattered to grouped, immersed to erumpent, globose
to subglobose in surface view, depressed globose to hemi-
spherical in section. Ostiolar neck or short papillate, central,
sometimes compressed, composed of heavily melanised cells.
Ascomatal wall composed of pale brown cells, with or without
“rim-like”stromatic region. Pseudoparaphyses septate, branched
and anastomosed. Asci fissitunicate, clavate to broadly cylin-
drical, stipitate, with 8 ascospores. Ascospores clavate to fusi-
form, with a supramedian primary septum, multiseptate, slightly
constricted at septa, hyaline, smooth, with an entire sheath.
Asexual morph where known sporodochial, monodictys-like
hyphomycetes.
Type genus:Parabambusicola Kaz. Tanaka & K. Hiray.
Notes: This family superficially resembles Bambusicola, but can
be distinguished from the latter by the ascomata surrounded by
stromatic tissue (Parabambusicola) or compressed necks with
wide ostioles (Aquastroma), and the multiseptate, clavate to
fusiform, hyaline ascospores.
Aquastroma Kaz. Tanaka & K. Hiray., gen. nov. MycoBank
MB811325.
Etymology: From the Latin aqua meaning water and stroma.
Ascomata scattered to grouped, immersed to erumpent, globose
in surface view, depressed globose in section. Ostiolar neck
central, compressed, composed of thick-walled, heavily melan-
ised, subglobose cells, with a wide ostiole. Ascomatal wall
composed of pale brown, thin-walled cells. Pseudoparaphyses
septate, branched and anastomosed. Asci fissitunicate, clavate,
stipitate, with 8 ascospores. Ascospores clavate to fusiform,
slightly curved, with a supramedian primary septum, multi-
septate, slightly constricted at septa, hyaline, smooth, with an
entire sheath. Asexual morph unknown.
Type species:Aquastroma magniostiolata Kaz. Tanaka &
K. Hiray.
Notes:Aquastroma is superficially similar to Quintaria in having
multiseptate ascospores and aquatic habitat. However, the type
species of Quintaria,Q. lignatilis known from marine habitats,
has long-necked ascomata with black incrustations surrounding
the sides of the ostiolar canal (Zhang et al. 2012), and is
phylogenetically close to members of the family Amniculicola-
ceae (Shearer et al. 2009) or the Testudinaceae/Platystomaceae
(Suetrong et al. 2009, Zhang et al. 2012). Quintaria is regarded
as a polyphyletic genus, because Q. submersa known from
freshwater habitats does not cluster with Q. lignatilis and forms a
sister clade to the Lindgomycetaceae (Zhang et al. 2012). An
asexual morph was not observed for Aquastroma in culture, but
this genus forms a clade with two hyphomycetes, Monodictys
species, and their relationship is highly supported (100 %;
Fig. 1). Monodictys, however, is not monophyletic and species in
this genus are phylogenetically distributed in three ascomycete
classes, Dothideomycetes,Leotiomycetes, and Sordariomycetes
(see Notes in Monodictys). Aquastroma should be retained as a
distinct genus, until sequence data of the type species of Mon-
odictys (M. putredinis) become available to clarify the phyloge-
netic placement of Monodictys s. str.
Aquastroma magniostiolata Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811326. Fig. 40.
Etymology: From the Latin mangi-, meaning large, and ostiola-
tus, meaning ostiolate.
Ascomata scattered to 3–4 grouped, immersed to erumpent,
globose in surface view, depressed globose in section,
180–250 μm high, 260–340 μm diam. Ostiolar neck central,
compressed, 60–750 μm long, 75–130 μm wide, composed of
thick-walled, heavily melanised, subglobose cells (2–3μm
diam), with a wide ostiole. Ascomatal wall in longitudinal section
25–30 μm thick at sides, composed of 5–7 layers of polygonal,
thin-walled, 6–17 × 2.5–7.5 μm, pale brown cells; wall at the
base 10–18 μm thick or poorly developed, composed of thin-
walled flattened cells. Pseudoparaphyses septate, branched
and anastomosed, 2–3μm wide. Asci fissitunicate, clavate,
112.5–137.5 × 25–29.5 μm, with a stipe of 22 –28 μm long,
with 8 biseriate to triseriate ascospores. Ascospores clavate to
fusiform, slightly curved, 30–44 × 9–13.5 μm (av.
36.5 × 10.3 μm, n = 39), l/w 2.8–4.5 (av. 3.6, n = 39), with a
supramedian primary septum (0.40–0.47; av. 0.43, n = 39),
6–8-septate (2+1+3, 2+1+4, 3+1+3, 3+1+4), slightly constricted
at septa, hyaline, smooth, with an entire sheath; sheath
delimited and 2–3μmwideatfirst, diffuse and up to 10 μm
wide at a later.
Colonies on PDA (after 4 wk) attaining a diam of 3.2–3.4 cm,
greenish grey; reverse almost black; no pigment produced. In
culture sexual morph formed. Asci and ascospores larger than
those on the natural host. Asci 130–170 × 22.5–37.5 μm (av.
148.9 × 29.9 μm, n = 30). Ascospores 39–47.5 × 11–16 μm (av.
43.5 × 13.5 μm, n = 50), l/w 2.8–3.8 (av. 3.2, n = 50).
Specimen examined:Japan, Mie, Tsu, Kamihama, Hyourtan pond, on submerged
twigs of woody plant, 1 Jun. 2008, K. Tanaka & H. Yonezawa (holotype KT
2485 = HHUF 30122, culture ex-type CBS 139680 = JCM 19429 = MAFF 243824).
Notes:Aquastroma magniostiolata is shown to be morphologi-
cally distinct from Quintaria aquatica by its ascospores with 6–8
septa [vs. (10–)11–13(–14) septa in Q. aquatica;Hyde & Goh
1999]. It is also different from Q. microspora in its larger asco-
spores (vs.26–31 × 5–5.5 μminQ. microspora;Zhang et al.
2008a).
Parabambusicola Kaz. Tanaka & K. Hiray., gen. nov. Myco-
Bank MB811327.
Etymology: After its phylogenetic similarity to Bambusicola.
Ascomata mostly grouped, immersed to erumpent, subglobose
in surface view, hemispherical with flattened base in section.
Ostiolar neck short papillate, central. Ascomatal wall at the sides
composed of pale brown, flattened cells; at the rim, composed of
parallel rows of rectangular to polygonal cells. Pseudopar-
aphyses numerous. Asci broadly cylindrical to clavate, short-
stalked, 8-spored. Ascospores fusiform, multiseptate, primary
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 115
septum mostly supramedian, hyaline, smooth, with an entire
sheath. Asexual morph unknown.
Type species:Parabambusicola bambusina (Teng) Kaz. Tanaka
& K. Hiray.
Notes: A new genus Parabambusicola is established to
accommodate P. bambusina formerly classified in Massarina
(Teng 1936, Aptroot 1998, Tanaka & Harada 2003b). Para-
bambusicola differs from Massarina in its hemispherical to
depressed globose ascomata surrounded by stromatic tissue
and without prominent clypeus, broadly cylindrical asci, and
fusiform ascospores with several septa. It is distantly related to
Massarina (Fig. 1).
Parabambusicola bambusina (Teng) Kaz. Tanaka & K. Hiray.,
comb. nov. MycoBank MB811392. Fig. 41.
Basionym:Massarina bambusina Teng, Sinensia 7: 512. 1936.
Specimens examined:Japan, Hokkaido, Chitose, Shikotsuko, on dead twigs of
Sasa kurilensis, 5 Sep. 2001, Y. Harada, H 4321 = HHUF 26590, culture MAFF
239462; Iwate, Hachimantai, Aspite line, on dead twigs of Sasa kurilensis, 25 Jul.
2009, K. Tanaka & Y. Harada, KT 2637 = HHUF 30120, culture MAFF 243822;
Aomori, Hirosaki, Tokiwano, Kuromori, Top of Mt. Iwaki, on dead twigs of Sasa
sp., 21 Jun. 2008, K. Hirayama & K. Tanaka, KH 139 = HHUF 30121, culture
MAFF 243823.
Note: For further information on this species, see Teng (1936)
and Tanaka & Harada (2003b).
Periconiaceae (Sacc.) Nann., Repert. mic. uomo 4: 482. 1934.
emend.
=Periconieae Sacc., Syll. Fung. 4: 235. 1886.
Ascomata scattered to grouped, immersed to erumpent, globose.
Ostiolar neck central, papillate, with hyaline periphyses. Asco-
matal wall in longitudinal section composed of several layers of
thin-walled, pale brown to brown cells. Pseudoparaphyses
cellular, branched, anastomosed. Asci fissitunicate, oblong to
cylindrical, 8-spored. Ascospores broadly fusiform, 1-septate,
hyaline, smooth, with an entire sheath. Conidiomata periconia- or
noosia-like. Conidiophores macronematous, mononematous,
sometimes lacking. Branches pale brown to brown, smooth to
slightly echinulate. Conidial heads spherical. Conidiogenous
cells monoblastic to polyblastic, discrete on stipe and branches.
Conidia globose to ellipsoidal, aseptate, catenate, brown, ver-
ruculose to echinulate.
Type genus:Periconia Tode.
Note: Although Periconiaceae (Nannizzi 1934) has long been
ignored in modern fungal systematics, it should be regarded as a
natural taxon in the suborder Massarineae.
AB
CE
DG
F
HJKLMIN
Fig. 40. Aquastroma magniostiolata. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F, G. Asci; H–M.
Ascospores (arrowheads indicate gelatinous sheath; J. in Black-Blue ink; K–M. in India ink); N. Germinating ascospore. A –E, G –J, N from KT 2485; F, K–M from culture KT
2485. Scale bars: A = 1 mm; B = 500 μm; C = 100 μm; D–N=10μm.
TANAKA ET AL.
116
Periconia Tode, Fung. mecklenb. sel. (Lüneburg) 2: 2. 1791.
Type species:Periconia lichenoides Tode.
Notes: Among more than 185 taxa described as Periconia
(http://www.indexfungorum.org, Aug. 2015), only two species
have been reported to have sexual morphs. They are P. igniaria
(the sexual morph was referred to as Didymosphaeria or
Massarina in the Dothideomycetes;Booth 1968, Aptroot 1998)
and P. prolifica (Remispora or Okeanomyces in the Sordar-
iomycetes;Kohlmeyer 1969, Pang et al. 2004), and therefore it
is obvious that the genus Periconia is not monophyletic. DNA
sequence data of the type species of this genus, P. lichenoides,
are currently unavailable, but we judged that the lineage of
Periconia in the Dothideomycetes, including typical members of
Periconia, such as P. byssoides,P. cookei,P. igniaria, and
P. digitata, corresponds to Periconia s. str., based on their
morphological similarities with P. lichenoides (Mason & Ellis
1953). In contrast, P. prolifica, a marine fungus in the Sordar-
iomycetes, does not have macronematous conidiophores
(Anastasiou 1963, Kohlmeyer 1969), and is not regarded as
Periconia s. str.
Periconia homothallica Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811328. Fig. 42.
Etymology: Referring to its homothallism.
Ascomata scattered, immersed to erumpent, globose,
140–190 μm high, 160–180 μm diam, with an ostiole. Ostiolar
neck central, papillate, 24–40 μm long, 37–62 μmwide,with
hyaline periphyses. Ascomatal wall in longitudinal section
uniformly 11–15 μm thick, composed of 4–6layersof
polygonal, thin-walled, 3–15 × 2–5μm, pale brown cells.
Pseudoparaphyses cellular, 2 –2.5 μm wide at the apex, up to
5μm wide at the base, branched, anastomosed. Asci fissitu-
nicate, cylindrical to lageniform, 85–119.5 × 13 –17.5 μm(av.
96.5 × 15.3 μm, n = 20), with a shallow ocular chamber, short-
stalked (3.5–6μm long), with 8 biseriate ascospores. Asco-
spores broadly fusiform, 22–31 × 7–10 μm (av. 26.3 × 8.7 μm,
n=60),l/w2.6–3.7 (av. 3.0, n = 60), with a nearly median
septum (0.48–0.53; av. 0.51, n = 38), hyaline, smooth, with an
entire sheath; sheath gelatinous, up to 10 μmwidewhenfresh,
later 1–2μmwide.
Colonies on PDA (after 2 wk) attaining a diam of 4.3–5.6 cm,
straw; reverse similar; no pigment produced. In culture sexual
morph formed. Ascospores slightly smaller than those on the
natural host, 20–28 × 6–9μm (av. 24.0 × 7.8 μm, n = 45), l/w
2.7–3.5 (av. 3.1, n = 45).
Specimen examined:Japan, Yamagata, Mogamigun, Sakekawa, Magarikawa-
river, riverbank, on dead leaves of Phragmites japonica, 13 Oct. 2002, Y. Ooki
& Y. Harada (holotype KT 916 = HHUF 29105, culture ex-type CBS
139698 = JCM 13100 = MAFF 239610).
Notes: Although we were not able to observe a typical periconia-
like asexual morph from our specimen and isolate, we introduce
this as a new species of Periconia based on the morphology of
the sexual morph. The above material shares many character-
istics with sexual species in Periconia s. str. (i.e., P. igniaria and
P. pseudodigitata), such as those with immersed globose
ascomata with papillate necks, a peridium composed of thin-
walled polygonal cells, cylindrical asci, and broadly fusiform
hyaline ascospores with nearly median septum. Periconia
homothallica, however, has shorter but wider asci than those of
P. igniaria (85–119.5 × 13–17.5 μmvs. 150–165 × 8–12 μm,
Booth 1968) and has wider ascospores than those of
P. pseudodigitata [22–31 × 7–10 μmvs. 19.5 –27( –32) × 5 –7
μm].
In our phylogenetic tree (Fig. 1), P. homothallica formed an
unsupported clade with P. igniaria, as well as species in other
hyphomycetous genera, such as “Sporidesmium”tengii that have
macronematous conidiophores and multiseptate conidia (Wu &
Zhuang 2005), and Noosia banksiae, a species that lacks con-
spicuous conidiophores (Crous et al. 2011a). The presence of
these genera in this lineage may indicate that Periconia should
be subdivided into several morphologically similar genera or that
the taxonomic status of these Noosia/Sporidesmium isolates
should be re-evaluated.
FGK
D
AB C
J
H
I
E
Fig. 41. Parabambusicola bambusina. A, B. Ascomata on the natural host surface; C. Ascomata in culture (on rice straw); D, E. Ascomata in longitudinal section; F. Ascomatal
wall; G. Pseudoparaphyses; H. Ascus; I, J. Ascospores (arrowheads indicate gelatinous sheath); K. Spermatia. A, B from KH 139; C, G–J from culture KH 139; D –F from KT
2637; K from culture KT 2637. Scale bars: A = 1 mm; B, C = 500 μm; D, E = 100 μm; F–K=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 117
AB
DF
I
K
C
EG
H
J
Fig. 42. Periconia homothallica. A, B. Ascomata on the natural host surface; C. Ascoma in culture (on rice straw); D. Ascoma in longitudinal section (in lactophenol cotton blue);
E. Ascomatal wall; F. Pseudoparaphyses; G. Ascus; H–J. Ascospores (arrowheads indicate gelatinous sheath; J. in India ink); K. Germinating ascospores. A, B, D –H, K from
KT 916; C, I, J from culture KT 916. Scale bars: A = 2 mm; B, C = 500 μm; D = 50 μm; E–K=10μm.
BA
L
E
MQ
GH
K
FJ
C
P
NO
D
I
Fig. 43. Periconia pseudodigitata. A, B. Ascomata on the natural host surface; C. Ascomata in longitudinal section (in lactophenol cotton blue); D. Ascomatal wall; E.
Pseudoparaphyses; F. Ascus; G–I. Ascospores (arrowheads indicate gelatinous sheath; H. in Black-Blue ink); J. Germinating ascospore; K. Conidiomata in culture (on rice
straw); L–O. Conidial heads and conidiophores; P, Q. Conidia. A, B, E, G–J from KT 1395; C, D, F from KT 644; K –M from culture KT 1195; N from culture KT 644; O–Q from
culture KT 1395. Scale bars: A = 1 mm; B, K = 500 μm; C = 100 μm; D–J, P, Q = 10 μm; L–O=20μm.
TANAKA ET AL.
118
Periconia pseudodigitata Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811329. Fig. 43.
Etymology: After its morphological and phylogenetic similarity to
Periconia digitata.
Ascomata numerous, scattered or 2–3 grouped, immersed to
erumpent, globose, 160–200 μm high, 130–250 μm diam.
Ostiolar neck central, papillate, 45–55 μm long, 45–65 μm wide,
with hyaline periphyses, surrounded by clypeus-like structure
composed of subglobose to polygonal, slightly thickened,
5–7×3–4μm, dark brown cells. Ascomatal wall in longitudinal
section 8–13 μm thick at side, 5–8μm thick at the base,
composed of 3–5 layers of thin-walled, 6–13 × 2–5μm, brown
cells. Pseudoparaphyses cellular, numerous, 2–2.5 μm wide at
the apex, 4–6μm wide at the base, septate, branched, anas-
tomosed, guttulate, associated with gelatinous material. Asci
fissitunicate, cylindrical, 70–110 × 10.5–15.5 μm (av.
88.4 × 12.2 μm, n = 33), rounded at the apex and with an apical
chamber, short-stalked (5–15 μm long), with 8 irregularly
biseriate ascospores. Ascospores broadly fusiform with rounded
ends, straight or slightly curved, 19.5–27(–32) × 5–7μm (av.
22.5 × 6.1 μm, n = 134), l/w 2.9–4.5 (av. 3.7, n = 134), with
almost median septum (0.48–0.55, av. 5.1, n = 36), slightly
constricted at the septum, hyaline, with or without guttules,
smooth, with an entire sheath; sheath gelatinous, 1–2μm wide
at side and 2–4μm wide at both ends in fresh, becoming
delimited sheath in dry condition. Senescent spores brown,
echinulate, 1-septate. Ascospores germinating from one or both
ends.
Colonies on PDA (after 4 wk) attaining a diam of 3.2–4.6 cm,
straw to amber with white margin; reverse pale luteous; no
pigment produced. In culture conidial and spermatial morphs
formed. Conidiophores macronematous, mononematous, sin-
gle or rarely 2–3 together on stromata, 250–450 μm long,
mostly 8.5–15 μmwide(16–24 μmwideatbasalstromatic
portion), 4–7-septate at 35–90 μm long intervals, with several
branches. Branches 10–50 μm long, septate, pale brown to
brown, slightly echinulate. Conidial heads spherical, ca.
60–120(–150) μmdiam.Conidiogenous cells monoblastic,
discrete on stipe and branches. Conidia catenate, globose,
brown, verruculose to shortly echinulate, 7–9.5 μm(av.8.3μm,
n=90).Spermogonia globose to subglobose, single to
gregarious, uniloculate to multiloculate, 140–340 μmdiam.
Spermatia subglobose, hyaline, smooth, 3–5×1.5–2μm(av.
3.7 × 1.7 μm, n = 30).
Specimens examined:Japan, Aomori, Hirakawa, Hirakawa-river, riverbank, on
dead culms of Phragmites australis, 5 Aug. 2001, K. Tanaka, KT 644 = HHUF
27569, culture JCM 13164 = MAFF 239674; Aomori, Hirosaki, Kawai, Hirakawa-
river, riverbank, 18 Aug. 2001, K. Tanaka, KT 680 = HHUF 27570; Aomori,
Hirosaki, Shimizumori, Oowasawa-river, riverbank, 14 Aug 2001, K. Tanaka, KT
673 = HHUF 27571; Hokkaido, Akkeshi, Toraibetsu-river, on submerged stems of
herbaceous plant, 3 Jun. 2003, K. Tanaka & S. Hatakeyama, KT 1195A = HHUF
29368, culture JCM 13165 = MAFF 239675; ibid., on dead culms of Phragmites
australis, 7 Sep. 2003, K. Tanaka & S. Hatakeyama (holotype KT 1395 = HHUF
29370, culture ex-type CBS 139699 = JCM 13166 = MAFF 239676); ibid., KT
1396–1398 = HHUF 29371–29373.
Notes:Periconia pseudodigitata is phylogenetically and
morphologically closest to P. digitata, but the conidial dimensions
of P. pseudodigitata (7 –9.5 μm) are slightly smaller than those
reported for P. digitata by Ellis (1971; 7 –11 μm). Sequence
similarity between these two taxa in the ITS region was 95.8 %
(503/525) with 0.8 % (4/525) gaps. Periconia pseudodigitata
superficially resembles P. igniaria (= Massarina igniaria), which
also has an ascomatal morph. The sexual morph of
P. pseudodigitata, however, differs from that of P. igniaria (Booth
1968) by its narrower ascospores (5–7μmvs.8–9μm) sur-
rounded by a mucilaginous sheath, and shorter but wider asci
(70–110 × 10.5–15.5 μmvs. 150 –165 × 8–12 μm) with biseriate
ascospores. The asexual morph of P. pseudodigitata has ver-
ruculose to shortly echinulate conidia, while that of P. igniaria has
conidia with conspicuous spines about 1 μm long (Mason & Ellis
1953, Ellis 1971, Matsushima 1971, Bunning & Griffiths 1984).
Sulcatisporaceae Kaz. Tanaka & K. Hiray., fam. nov. Myco-
Bank MB814431.
Etymology: Referring to the name of the type genus.
Ascomata scattered to grouped, immersed to erumpent, globose
in surface view, subglobose to hemispherical in section. Ostiolar
neck short papillate, central, with periphyses. Ascomatal wall
composed of several layers of compressed cells, poorly devel-
oped at the base. Pseudoparaphyses trabeculate, branched and
anastomosed. Asci clavate, short-stalked, 8-spored. Ascospores
broadly fusiform, 1-septate, hyaline, with an entire sheath.
Conidiomata pycnidial, globose. Conidiogenous cells cylindrical
to doliiform, annellidic or phialidic. Conidia ellipsoid to sub-
globose, hyaline to dark brown, 1- to multi-septate (occasionally
muriform), with or without striate ornamentation.
Type genus:Sulcatispora Kaz. Tanaka & K. Hiray.
Note: A new family, Sulcatisporaceae is established here for
Magnicamarosporium and Sulcatispora genera nova and Neo-
bambusicola (Crous et al. 2014b).
Magnicamarosporium Kaz. Tanaka & K. Hiray., gen. nov.
MycoBank MB811292.
Etymology: After its morphological similarity to Camarosporium
and its large-sized conidiomata.
Conidiomata pycnidial, scattered, immersed, depressed globose
in section. Ostiolar neck central, papillate to short cylindrical.
Paraphyses filamentous, sometimes branched, associated with
gelatinous material. Conidiophores absent. Conidiogenous cells
holoblastic, cylindrical to doliiform. Conidia subglobose to
obovoid, sometimes obpyriform, muriform, dark brown. Sexual
morph unknown.
Type species:Magnicamarosporium iriomotense Kaz. Tanaka &
K. Hiray.
Notes: Several coelomycetous genera are known to have dic-
tyosporous pigmented conidia and their taxonomic revision has
been reported (e.g., Van Warmelo & Sutton 1981, Nag Raj 1989,
Verkley et al. 2005, Crous et al. 2015b). Amarenographium is
characterised by longer and branched conidiophores and conidia
with cap-like gelatinous appendages (Eriksson 1982). Species in
this genus are reported from marine environments (Kohlmeyer &
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 119
Volkmann-Kohlmeyer 1991) or palms (Taylor & Hyde 2003), and
may be genetically close to the Trematosphaeriaceae (Hodhod
et al. 2012). Camarosporellum and Camarographium have dis-
toseptate conidia (Sutton 1980, Verkley et al. 2005) rather than
euseptate conidia like Magnicamarosporium.Camarographium
is heterogenous, and C. koreanum belongs to Unknown Clade III
of the Massarineae (Fig. 1), while C. carpini may have affinity
with the Sporormiaceae based on a BLAST search using LSU
and ITS nrDNA sequences (Crous et al. 2011b). Camar-
osporium, to some degree resembles our new genus, but
Magnicamarosporium has larger conidiomata with a conspicuous
projecting ostiole composed of thick-walled cells, long paraphy-
ses with a gelatinous coating, and larger dark brown conidia than
Camarosporium. In addition, Camarosporium does not have
phylogenetic affinity with Magnicamarosporium, because the
type species of Camarosporium (C. quaternatum) is a member of
the Pleosporineae, based on molecular data (Crous et al. 2006,
Wijayawardene et al. 2014a, b, d, Crous et al. 2015b).
The phylogenetic tree generated in this study indicated that
Magnicamarosporium and Neobambusicola cluster together with
74 % BP (Fig. 1). However, the conidial morphology of Neo-
bambusicola (fusoid-ellipsoid and 1-septate; Crous et al. 2014b)
is quite distinct from that of Magnicamarosporium.
Magnicamarosporium iriomotense Kaz. Tanaka & K. Hiray.,
sp. nov. MycoBank MB811293. Fig. 44.
Etymology: Referring to the location where the specimen was
collected.
Conidiomata pycnidial, scattered, deeply immersed, depressed
globose in section, 330 –440 μm high, 700–760 μm diam. Ostiolar
neck central, papillate to short cylindrical, 120–150 μm long,
80–100 μm wide, composed of polygonal to rectangular, thick-
walled, dark brown cells of 2–5μm diam. Conidiomatal wall in
longitudinal section uniformly 10–20 μm thick, composed of 3–6
layers of polygonal to subglobose, thin-walled, pale brown cells of
2–7.5 × 2–5μm. Paraphyses sometimes branched,
20–50(–80) μm long, 1.5–2.5 μm wide, associated with gelati-
nous material. Conidiophores absent. Conidiogenous cells holo-
blastic, 7–11 × 4.5–6μm, cylindrical to doliiform. Conidia
subglobose to obovoid, sometimes obpyriform, 29 –40.5(–42.5) ×
19.5–25(–27) μm (av. 34.7 × 22.2 μm, n = 100), l/w 1.2–2.0 (av.
1.6, n = 100), with 4–6 trans- and 1–2 vertical-septa, frequently
with oblique septa, dark brown, smooth, without sheath.
Colonies on PDA (after 4 wk) attaining a diam of 1.1–1.5 cm,
olivaceous black with buff margin; reverse buff to honey; no
pigment produced. In culture asexual morph formed. Con-
idiomata pycnidial, 130–170 μm diam, scattered to 2–3 grouped,
globose to subglobose. Conidia 29.5–43 × 18–23 μm (av.
35.3 × 20.3 μm, n = 31), l/w 1.5–2.1 (av. 1.7, n = 31). Sexual
morph unknown.
Specimen examined:Japan, Okinawa, Isl. Iriomote, Tropical botanic garden, on
dead twigs of Diplospora dubia, 13 Jul. 2011, K. Tanaka & K. Hirayama (ho lotype KT
2822 = HHUF 30125, culture ex-type CBS 139696 = JCM 19402 = MAFF 243827).
Notes: There are more than 500 names in Camarosporium
(http://www.indexfungorum.org, Aug. 2015). Many of these taxa
have been described on the basis of their host associations
(Sutton 1980, Marincowitz et al. 2008), but there is no record of
Camarosporium from host plants within the Rubiales, which in-
cludes Diplospora dubia, the host of M. iriomotense.
Sulcatispora Kaz. Tanaka & K. Hiray., gen. nov. MycoBank
MB811294.
Etymology: From the Latin sulcatus meaning “furrowed”and
spora meaning “spore”, referring to the striate ornamentation of
the conidia.
Ascomata scattered to grouped, immersed to erumpent, globose
in surface view, subglobose to hemispherical in section. Ostiolar
neck short papillate, central, with periphyses. Ascomatal wall
composed of several layers of compressed cells, poorly devel-
oped at the base. Pseudoparaphyses trabeculate, branched and
anastomosed. Asci fissitunicate, clavate, short-stalked. Asco-
spores broadly fusiform, 1-septate, hyaline, with an entire
sheath. Conidiomata pycnidial, globose. Conidiogenous cells
cylindrical, annellidic. Conidia ellipsoid, yellowish brown, multi-
septate, with striate ornamentation.
Type species:Sulcatispora acerina Kaz. Tanaka & K. Hiray.
Notes: Ascomatal features of Sulcatispora are most similar to
those of Massarina s. str. (Massarinaceae) but the ascomata lack
a prominent clypeus, in contrast to species within Massarina s.
str. The most distinctive feature of Sulcatispora is the striated
conidia. Some species in Phaeophleospora (e.g., P. striae;Taylor
& Hyde 2003) are known to have such conidia, but Phaeo-
phleospora (type species: P. eugeniae) is phylogenetically placed
in the Mycosphaerellaceae of the Capnodiales (Crous et al.
2007, 2009b). Sclerostagonospora also has species with con-
idia with striate ornamentation (e.g., S. opuntiae;Huhndorf
1992), but species of Sclerostagonospora have phylogenetic
affinities with members of the Phaeosphaeriaceae, such as
“Phoma”caloplacae and “Phoma”foliaceiphila (Crous et al.
2011a, Lawrey et al. 2012), and may have phaeosphaeria-like
sexual morphs (Quaedvlieg et al. 2013). Sulcatispora is phylo-
genetically related to Bambusicola, which also has a similar
asexual morph. Bambusicola, however, has conical ascomata
with flattened bases, narrower asci (up to 14 μm wide), and
narrowly fusiform ascospores (Dai et al. 2012).
Sulcatispora acerina Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811295. Fig. 45.
Etymology: Referring to the generic name of the host.
Ascomata scattered, immersed to erumpent, globose in surface
view, depressed globose to hemispherical in section,
200–250 μm high, 260–450 μm diam. Ostiolar neck incon-
spicuous short papillate, 60–85 μm diam, central, with periph-
yses. Ascomatal wall in longitudinal section 12–25 μm thick at
sides, composed of polygonal, 5–8×2–3μm, compressed cells,
surrounded by short, brown hyphae (2.5–3.5 μm thick), poorly
developed at the base. Pseudoparaphyses trabeculate,
1.5–2μm wide, branched and anastomosed, associated with
gelatinous material. Asci fissitunicate, clavate,
87–113 × 15–20 μm (av. 98.4 × 17.1 μm, n = 17), rounded at the
apex, with an apical chamber and faint ring, short-stalked
(15–22 μm long). Ascospores broadly fusiform, slightly acute
at the apex, (23–)26–32 × 7–9μm (av. 28.5 × 7.7 μm, n = 50), l/
TANAKA ET AL.
120
w 3.2–4.2 (av. 3.7, n = 50), with a submedian primary septum
(0.50–0.57; av. 0.53, n = 50), hyaline, surrounded by an entire
sheath; sheath 2–3μm wide at first, later diffuse, 4–5μm wide.
Colonies on PDA (after 4 wk) attaining a diam of 2.1–2.4 cm,
grey olivaceous to honey; reverse olivaceous to black; no
pigment produced. In culture asexual morph formed. Con-
idiomata pycnidial, 180–270 μm high, 160–240 μm diam,
immersed, scattered, globose, ostiolate. Ostiolar neck short
papillate, 25–45 μm long, 35–45 μm diam. Conidiomatal wall
uniformly 10–20 μm thick, composed of 4–7 layers of polygonal,
6–10 × 2–3μm, dark brown cells. Conidiophores absent.
Conidiogenous cells cylindrical to lageniform, 10–20 μm long,
2μm wide at the apex, 3.5–5.5 μm wide at the base, annellidic.
Conidia ellipsoid, rounded at the apex, truncate at the base,
yellowish brown, 20–28(–30) × 6.5–8μm (av. 25.9 × 7.2 μm,
n = 50), l/w 3.1–4.2 (av. 3.6, n = 50), 3(–5)-septate (1+1+1,
rarely 2+1+2), with striate ornamentation.
Specimen examined:Japan, Fukuoka, Kasuya, Hisayama, Yamada, Mt. Tachi-
banayama, on dead twigs of Acer palmatum, 31 Mar. 2012, K. Tanaka (holotype
KT 2982 = HHUF 30449, culture ex-type CBS 139703).
Note: The conidia of S. acerina are very similar to those of
S. berchemiae, but S. acerina can be distinguished from the
latter by its larger ascospores [(23–)26–32 × 7–9μmvs.
22–29 × 5.5–7μm].
Sulcatispora berchemiae Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811296. Fig. 46.
Etymology: Referring to the generic name of the host.
Ascomata scattered to 2–4 grouped, immersed to erumpent,
globose in surface view, subglobose to hemispherical in section,
150–280 μm high, 350–530 μmdiam.Ostiolar neck short papillate,
ca. 60 μm diam, central, with periphyses. Ascomatal wall in longi-
tudinal section 10–18 μm thick at sides, composed of 3 –6layersof
7–18 × 2 μm compressed cells, poorly developed at the base.
Pseudoparaphyses trabeculate, 1.5–2.5 μm wide, branched and
anastomosed, associated with gelatinous material. Asci fissituni-
cate, clavate, 77.5–100 × 13.5 –18 μm (av. 90.1 × 15.2 μm, n = 20),
rounded at the apex, with an apical chamber and faint ring, short-
stalked (13–15 μmlong).Ascospores broadly fusiform,
22–29 × 5.5–7μm (av. 26.0 × 6.4 μm, n = 50), l/w 3.5–4.4(–4.7)
(av. 4.1, n = 50), with a submedian primary septum (0.50–0.57; av.
0.53, n = 49), hyaline, surrounded by an entire sheath; sheath
delimited, 2 μmwideatfirst, later diffuse, up to 6 μmwide.
Colonies on PDA (after 4 wk) attaining a diam of 2.7–2.8 cm,
white to buff; reverse buff to greyish sepia; no pigment produced.
In culture asexual morph formed. Conidiomata pycnidial,
90–130 μm high, 90–150 μm diam, immersed, scattered,
globose to subglobose, ostiolate. Ostiolar neck short papillate.
Conidiomatal wall uniformly 12–17.5 μm thick, composed of
flattened, thin-walled, polygonal cells (3.5–6 × 1.5–2.5 μm).
Conidiophores absent. Conidiogenous cells cylindrical,
12–15 μm long, 2 μm wide at the apex, 4–8μm wide at the
base, annellidic. Conidia ellipsoid, 20–30 × 6.5–8μm (av.
24.8 × 7.1 μm, n = 50), l/w 2.9–4.2 (av. 3.5, n = 50), rounded at
B
F
C
H
ML
K
N
JI
G
DE
A
Fig. 44. Magnicamarosporium iriomotense. A. Immersed conidiomata and discharged conidia lying on the natural host surface; B. Conidiomata produced in culture (on rice
straw); C, D. Conidiomata in longitudinal section (C. on nature; D. in culture); E. Conidiomatal wall; F. Paraphyses; G. Conidiogenous cell (in trypan blue); H–L. Conidia; M.
Immature conidium; N. Germinating conidium. A, C, E, H–J, N from KT 2822; B, D, F, G, K–M from culture KT 2822. Scale bars: A, B = 500 μm; C, D = 50 μm; E –N=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 121
the apex, truncate at the base, yellowish brown, 3(–5)-septate
(1+1+1, rarely 2+1+2), with striate ornamentation.
Specimens examined:Japan, Aomori, Towada, Sanbongi, Yagami, on vines of
Berchemia racemosa, 2 Dec. 2003, K. Tanaka, S. Hatakeyama & N. Nakagawara
(holotype KT 1607 = HHUF 29097, culture ex-type CBS 139704 = JCM
13101 = MAFF 239611); ibid., KT 1608 = HHUF 29098.
Note: This species is closely related to S. acerina with regard to its
LSU sequences (similarity 1295/1302 = 99.5 %), but the ITS
sequence shows a great deal of variation compared to the ITS of
S. acerina (similarity 770/850 = 90.6 %, with gaps29/850 = 3.4 %).
Trematosphaeriaceae K.D. Hyde et al., Cryptog. Mycol. 32:
347. 2011.
A
B
DC
EF GHIJKL
Q
V
O
M
P
S
N
TUR
W
Fig. 45. Sulcatispora acerina. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G–K.
Ascospores (arrowheads indicate mucilaginous sheath); L. Germinating ascospore; M, N. Conidiomata in culture (on rice straw); O. Conidioma in longitudinal section; P.
Conidiomatal wall; Q. Conidiogenous cells (arrowheads indicate annellations); R–W. Conidia. A–L from KT 2982; M –W from culture KT 2982. Scale bars: A = 1 mm; B, M,
N = 500 μm; C, O = 50 μm; D–L, P–W=10μm.
TANAKA ET AL.
122
Type genus:Trematosphaeria Fuckel.
Trematosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24: 161.
1870.
Type species:Trematosphaeria pertusa Fuckel.
Notes: Although more than 200 species have been assigned to
Trematosphaeria (http://www.indexfungorum.org, Aug. 2015),
most of these species have not had their generic placements
verified by molecular evidence. The type species of this genus,
T. pertusa, usually grows on terrestrial wood, but also can survive
within freshwater (Suetrong et al. 2011b). Recently, Madurella
grisea, originating from a human infection, water and pastry gel,
was added to Trematosphaeria based on a multi-gene phylogeny
(Ahmed et al. 2014).
Trematosphaeria pertusa Fuckel, Jb. Nassau. Ver. Naturk.
23–24: 161. 1870. Fig. 47.
Ascomata gregarious, immersed or becoming superficial by
weathering of host tissue, globose to pyriform in section,
R
QP
O
N
M
E
L
BC
K
I
G
H
J
S
F
A
D
Fig. 46. Sulcatispora berchemiae. A, B. Ascomata on the natural host surface; C, D. Ascomata in longitudinal section; E. Ascomatal wall; F. Pseudoparaphyses; G. Ascus; H–J.
Ascospores (arrowheads indicate mucilaginous sheath); K. Germinating ascospore; L. Conidiomata in culture (on rice straw); M. Conidioma in longitudinal section; N. Con-
idiomatal wall; O. Conidiogenous cells (arrowheads indicate annellations); P–S. Conidia. A–K from KT 1607; L –S from culture KT 1607. Scale bars: A = 1 mm; B, L = 500 μm;
C, D = 100 μm; E–K, M–S=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 123
270–380 μm high, 190–350 μm diam. Ostiolar neck central,
papillate to cylindrical, 70–140 μm long, 90–130 μm wide,
composed of thick-walled, heavily melanised cells (2–4μm
diam), with periphyses. Ascomatal wall in longitudinal section
uniformly 17–25 μm thick, composed of 6–9 layers of polygonal
to rectangular, thin-walled, brown cells (2–15 × 2 –5μm).
Pseudoparaphyses branched and anastomosed, 1.5–2.5 μm
wide. Asci fissitunicate, clavate to cylindrical,
73.5–102.5 × 12.5–17 μm (av. 89.8 × 14.6 μm, n = 20), with a
stipe of 8.5–22 μm long, with 8 biseriate ascospores. Asco-
spores fusiform, slightly curved, 20–27.5 × 6–8μm (av.
23.9 × 7.1 μm, n = 50), l/w 2.8–4.0 (av. 3.4, n = 50), with a nearly
median or somewhat supramedian primary septum (0.43–0.53;
av. 0.48, n = 50), rarely 3-septate, slightly constricted at the
septum, reddish brown, smooth, without sheath.
In culture spermatial morph formed. Spermatia subglobose,
hyaline, 2.8–3.9 × 1.9–2.3 μm (av. 3.4 × 2.1 μm, n = 20), l/w
1.4–1.9 (av. 1.6, n = 20).
Specimens examined:Japan, Aomori, Hirosaki, Aoki, Mohei-pond, on sub-
merged twigs of woody plant, 27 Sep. 2003, K. Tanaka & N. Asama, KT
1496 = HHUF 30153, culture JCM 19430 = MAFF 243879; Hokkaido, Isl. Rebun,
Nairo, Nairo-river, on submerged twigs of woody plant, 16 Aug. 2013, K. Tanaka,
KT 3314 = HHUF 30452, culture CBS 139705; ibid., KT 3315 = HHUF 30453,
culture CBS 139706.
Notes: We identified our specimens/isolates as T. pertusa, the type
species of Trematosphaeria, based on close similarities between
sequences obtained from the above isolates and an ex-epitype
strain of the species (CBS 122368; Ahmed et al. 2014), i.e., 484/
485 (99.8 %) in ITS (GenBank KF015668) and 900/907 (99.2 %) in
tef1 (GenBank KF015701) regions. In comparison with the
description of T. pertusa based on the neotype (Zhang et al. 2008b),
our specimens haveshorter asci (73.5–102.5 μmvs.100–145 μm)
and ascospores (av. 20–27.5 μmvs.27.5–32.5 μm). Further
collections of this species are needed to clarify the taxonomic
significance of the intraspecific morphological variation.
Unknown Clade I
Fuscostagonospora Kaz. Tanaka & K. Hiray., gen. nov.
MycoBank MB811330.
Etymology: From the Latin fusco-, meaning dark brown and the
generic name, Stagonospora.
Ascomata scattered, immersed, globose to subglobose. Ostiolar
neck clypeate, central, short papillate, with periphyses. Asco-
matal wall composed of pale brown, compressed cells. Pseu-
doparaphyses appearing trabecular, branched and
anastomosed, associated with gelatinous material. Asci fissitu-
nicate, cylindrical, with a long stipe. Ascospores narrowly fusi-
form, 1(–3)-septate, hyaline, with an entire sheath. Conidiomata
pycnidial, scattered, immersed, depressed globose, ostiolate.
Conidiomatal wall composed of thin-walled cells. Conidiophores
absent. Conidiogenous cells doliiform, annellidic. Conidia yellow
to pale brown and 3-septate.
Type species:Fuscostagonospora sasae Kaz. Tanaka & K. Hiray.
Notes: The new genus, Fuscostagonospora, is introduced to
accommodate the bambusicolous fungus, F. sasae. This genus
is reminiscent of Stilbospora in having pigmented phragmospo-
rous conidia, but the latter genus has acervular conidiomata filled
with paraphyses and phylogenetically groups within the
L
J
I
H
G
EF
DC
B
A
MK
Fig. 47. Trematosphaeria pertusa. A, B. Ascomata on the natural host surface; C. Ascoma in longitudinal section; D. Ascomatal wall; E. Pseudoparaphyses; F. Ascus; G–I.
Ascospores; J. Germinating ascospore; K. Spermogonia in culture; L. Spermatiophores; M. Spermatia. A–J from KT 1496; K –M from culture KT 1496. Scale bars: A,
K = 500 μm; B = 200 μm; C = 50 μm; D–J, L, M = 10 μm.
TANAKA ET AL.
124
Diaporthales in the Sordariomycetes (Crous et al. 2012,
Voglmayr & Jaklitsch 2014). The conidial morphology of Fus-
costagonospora is also similar to that of Sclerostagonospora, but
the latter genus has phylogenetic affinity with species in the
Phaeosphaeriaceae (Pleosporales) and may have a
phaeosphaeria-like sexual morph (Quaedvlieg et al. 2013). In
overall morphology, such as the narrowly fusiform ascospores
and pigmented septate conidia, as well as in host preferences,
Fuscostagonospora is most similar to Bambusicola (Bambusi-
colaceae;Dai et al. 2012). Fuscostagonospora, however, has
ascomata with a prominent clypeus and phylogenetically de-
viates from the Bambusicolaceae, forming an Unknown Clade I
(Fig. 1).
Fuscostagonospora sasae Kaz. Tanaka & K. Hiray., sp. nov.
MycoBank MB811331. Fig. 48.
Etymology: Referring to the host genus.
Ascomata scattered, immersed, globose to subglobose,
220–250 μm high, 400–450 μmdiam.Ostiolar neck clypeate,
central, short papillate, 50–65 μmlong,75–90 μm wide, with
periphyses. Ascomatal wall in longitudinal section 10–15 μm thick
at sides, composed of 3–4 layers of compressed, 5–10 × 2 –3μm,
pale brown cells. Pseudoparaphyses appearing trabecular,
1–1.5 μm wide, branched and anastomosed, associated with
gelatinous material. Asci fissitunicate, cylindrical,
90–140 × 8.5 –14.5 μm (av. 107.6 × 9.8 μm, n = 30), with an apical
chamber, with a relatively long stipe (20–32.5 μm long). Asco-
spores narrowly fusiform, 1(–3)-septate, 22–31.5(–36) ×
3–5.5 μm (av. 27.5 × 4.1 μm, n = 25), l/w 5.6–8.3 (av. 6.8, n = 25),
with a nearly median septum (0.48–0.57; av. 0.52, n = 20), hyaline,
surrounded by an entire gelatinous sheath up to 7 μmwide.
D
N
B
E
FK
O
Q
P
R
C
GJ
IH
A
M
L
Fig. 48. Fuscostagonospora sasae. A, B. Ascomata on the natural host surface; C, D. Ascomata in longitudinal section; E. Ascomatal wall; F. Pseudoparaphyses; G. Ascus;
H–J. Ascospores (arrowheads indicate gelatinous sheath; H, I. in India ink); K. Germinating ascospore; L, M. Conidiomata in culture (on rice straw); N. Conidioma in longitudinal
section; O. Conidiomatal wall; P, Q. Conidiogenous cells (arrowheads indicate annellations); R. Conidia. A–K, N, O from KT 1467; L, M, P –R from culture KT 1467. Scale bars:
A, L = 1 mm; B, M = 500 μm; C, D, N = 100 μm; E–K, O–R=10μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 125
Conidiomata on the natural host pycnidial, scattered, immersed,
depressed globose in section, 250 μm high, 500 μm diam, ostiolate.
Conidiomatal wall in longitudinal section 12–20 μmthick,
composed of thin-walled, hyaline to pale brown cells. In culture
same coelomycetous asexual morph formed. Conidiophores ab-
sent. Conidiogenous cells doliiform, annellidic, 5–12 × 3 –7μm(av.
9.7 × 5.3 μm, n = 20). Conidia hyaline and aseptate at first, later
yellow to pale brown and 3-septate, 17–22.5 × 4–5μm(av.
19.8 × 4.4 μm, n = 50), l/w (3.4–)3.7–5.5 (av. 4.5, n = 50).
Colonies on PDA (after 4 wk) attaining a diam of 3.7–4.7 cm,
iron grey; reverse similar; no pigment produced. In culture
asexual morph formed.
Specimen examined:Japan, Fukushima. Minamiaizu, Ose pond, on dead twigs
of Sasa sp., 30 Aug. 2003, N. Asama (holotype KT 1467 = HHUF 29106, culture
ex-type CBS 139687 = JCM 13104 = MAFF 239614).
Unknown Clade IV
Pseudoxylomyces Kaz. Tanaka & K. Hiray., gen. nov. Myco-
Bank MB811332.
Etymology: After its morphological similarity to Xylomyces.
Colonies on natural substratum scattered, dark brown, glistening.
Mycelium immersed in agar medium, pale brown to reddish brown.
Stromata lacking. Conidiophores branched, septate, brown. Con-
idiogenesis holoblastic. Conidia broadly fusiform, with several
trans-septa of thick-walled, yellowish brown to dark brown with
paler end cells. Chlamydospores absent. Sexual morph unknown.
Type species:Pseudoxylomyces elegans (Goh et al.) Kaz.
Tanaka & K. Hiray.
Notes: A new genus Pseudoxylomyces is proposed to
accommodate Xylomyces elegans. The type species of Xylo-
myces (X. chlamydosporus) is known to have phylogenetic
affinities with the freshwater genus, Jahnula in the Jahnulales
(Campbell et al. 2007, Sivichai et al. 2011). Therefore, Xylo-
myces is now regarded as a synonym of Jahnula (Hyde et al.
2013). Xylomyces elegans was excluded from the genus
(Suetrong et al. 2011a), because it did not cluster with the type
species of Xylomyces in phylogenetic analyses using ITS
(Prihatini et al. 2008) and SSU sequences (Shearer et al.
2009). However, no taxonomic decision has been made for
X. elegans.
Xylomyces is characterised by its lack of conidiophores and
conidiogenous cells but it produces large, dark, thick-walled,
multiseptate, intercalary, narrowly fusiform chlamydospores
(Goos et al. 1977, Goh et al. 1997). In contrast, Pseudox-
ylomyces typified by P. elegans is quite distinctive in producing
broadly fusiform conidia holoblastically at the tip of the co-
nidiophores (Fig. 49A, B).
Pseudoxylomyces elegans (Goh et al.) Kaz. Tanaka & K.
Hiray., comb. nov. MycoBank MB811333. Fig. 49.
Basionym:Xylomyces elegans Goh et al., Mycol. Res. 101:
1324. 1997.
Mycelium immersed in agar, pale brown to reddish brown. Co-
nidiophores up to 40 μm long. Conidiogenesis holoblastic.
Conidia broadly fusiform, 4–7-septate, 72.5–98 × 35–41 μm, l/w
2.0–2.8, brown, with paler end cells. Sexual morph unknown.
Specimen examined:Japan, Okinawa, Isl. Iriomote, Oomijya-river, on sub-
merged twigs of woody plant, 12 Jul. 2011, K. Tanaka & K. Hirayama, KT
2887 = HHUF 30139, culture MAFF 243852.
Notes: This species appears to be widely distributed and has
been reported from Australia, Seychelles (Goh et al. 1997),
Thailand (Sivichai et al. 2000), Hong Kong (Tsui & Hyde 2004),
USA (Raja et al. 2007), Brazil (Barbosa & Gusm~
ao 2011), and
India (Patil & Borse 2015). A BLAST search using ITS se-
quences from our culture showed 98.7 % (464/470) similarity to
sequences of X. elegans (GenBank FJ887920) collected from
Thailand (Prihatini et al. 2008).
Unknown Clade VI
Monodictys S. Hughes, Canad. J. Bot. 36: 785. 1958.
Type species:Monodictys putredinis (Wallr.) S. Hughes, Canad.
J. Bot. 36: 785. 1958.
Notes: More than 60 species have been described in Monodictys
(http://www.indexfungorum.org, Aug. 2015), but their phylogenetic
placements are mostly unknown. Apparently, Monodictys is a
heterogenous group of hyphomycetes. Several species within this
genus such as M. arctica (Leptosphaeriaceae;Day et al. 2006), M.
cf. putredinis (Melanommataceae;Samuels 1980), Monodictys
spp. (Parabambusicolaceae;Fig. 1), and M. capensis (Unknown
Clade IV; Fig. 1) are scattered within the Dothideomycetes.Mon-
odictys pelagica is known to have phylogenetic relationships with
the Sordariomycetes (Mouzouras & Jones 1985, Campbell et al.
2002), and Monodictys sp. is reported as an asexual morph of
Hyaloscypha albohyalina var. monodictys (Hosoya & Huhtinen
A B
C F
E
D
Fig. 49. Pseudoxylomyces elegans. A, B. Conidiophores and conidia; C –E.
Conidia (E. bleached conidium); F. Germinating conidium. A, B, D, E from culture
KT 2887; C, F from KT 2887. Scale bars: A = 200 μm; B–F=20μm.
TANAKA ET AL.
126
2002;currentlyHyaloscypha monodictys,Han et al. 2014)inthe
Leotiomycetes. Phylogenetic reassessment of many species of
Monodictys including the type species of thisgenus (M. putredinis)
is needed for taxonomic revision of this genus.
Monodictys capensis R.C. Sinclair et al., Mycotaxon 59: 359.
1996. Fig. 50.
Specimen examined:Russia, St. Petersburg, Botanical garden of the Komarov
Botanical Institute of the Russian Academy of Sciences, on dead wood of Padus
avium, 19 Oct. 2005, V. Mel'nik, HR 1 = HHUF 29712 = LE 226298, culture CBS
134928 = VKM F-4506.
Notes: This material has been reported as M. capensis by Mel’nik
& Shabunin (2010). In culture, abundant conidia similar to those
on the natural host were produced on rice straw agar (RSA).
DISCUSSION
New familial lineages
The Parabambusicolaceae is erected here to accommodate
Aquastroma (Fig. 40) and Parabambusicola (Fig. 41), as well as
two unnamed “Monodictys”species. This family is superficially
similar to the Bambusicolaceae in having depressed globose to
hemispherical ascomata. In particular, Parabambusicola, the
type genus of the Parabambusicolaceae, has a bambusicolous
habitat like that of Bambusicola (Dai et al. 2012). However,
members of the Parabambusicolaceae have ascomata sur-
rounded by stromatic tissue (Parabambusicola) or compressed
necks with wide ostioles (Aquastroma), and also multiseptate,
clavate to fusiform, hyaline ascospores, unlike those of the
Bambusicolaceae. Asexual morphs possessing sporodochial
conidiomata and muriform conidia like those of Monodictys are
not found in the Bambusicolaceae.
A new family Sulcatisporaceae is introduced to encompas
Magnicamarosporium and Sulcatispora genera nova, as well as
Neobambusicola (Crous et al. 2014b). These three genera form
a strongly supported clade (97 %), which is a sister of the
Bambusicolaceae. The sexual morphs of species in the
Sulcatisporaceae are superficially similar to those of Bambusi-
colaceae, but subglobose to obovoid conidia with muriform
septation (Magnicamarosporium) or 1 to several septate conidia
with or without striate ornamentation (Neobambusicola,Sulca-
tispora) are not found in species of the Bambusicolaceae.
We resurrect the Periconiaceae as a sister taxon of the
Massarinaceae (Fig. 1). The name of “Periconieae”was origi-
nally used for dematiaceous hyphomycetes that have macro-
nematous conidiophores and 1-celled, pigmented conidia
forming conidial heads (Saccardo 1886). Later, the group was
raised to familial rank, and Periconia and Stachybotrys were
assigned to the Periconiaceae (Nannizzi 1934), the latter genus
now being placed in its own family, Stachybotryaceae (Crous
et al. 2014a). The Periconiaceae has long been ignored in
modern fungal systematics, but it should be regarded as a
natural taxon in the suborder Massarineae. The main genus
Periconia has been treated as a member of the Massarinaceae
based on a topology of genealogical trees (Kodsueb et al. 2007,
Schoch et al. 2009, Zhang et al. 2009b, c, 2012, Hyde et al.
2013). Species of Periconia, however, are separated from the
Massarinaceae and form a strongly supported clade of the
Periconiaceae (100 %; Fig. 1). Sexual morphs within the Per-
iconiaceae (Figs 42, 43) differ from those of Helminthosporium
(Fig. 25)andMassarina (Fig. 26)intheMassarinaceae.
Members in the latter family possess medium to large sub-
globose ascomata, which are covered by a clypeus, and have
clavate asci. The small ascomata of Periconia superficially
resemble those of Stagonospora (= Saccharicola,Massar-
inaceae;Figs 27–29) but differ in having a peridium composed
of small-sized compressed cells. The most diagnostic features
of the Periconiaceae are the asexual morphs of Periconia;
these have macronematous, mononematous conidiophores
with globose to cylindrical, blastic conidia (Mason & Ellis 1953,
Ellis 1971).
Dictyosporium and phenotypically similar genera, such as
Aquaticheirospora and Pseudodictyosporium, have been
considered to belong in the Massarinaceae (Wijayawardene
et al. 2012). However, the group including dictyosporium-like
fungi forms a distinct lineage (as Dictyosporiaceae nom. prov.;
see Liu et al. 2015), independent from the Massarinaceae
(Fig. 1). One of the diagnostic features of the Dictyosporiaceae is
their multicellular cheiroid conidia (Fig. 2), and these morpho-
logical features separate it from other families in the Massar-
ineae. Sexual morphs observed in the Dictyosporiaceae
(Gregarithecium and Pseudocoleophoma;Figs 3–5) are some-
what similar to those of the Massarinaceae (e.g., Stagonospora;
Figs 27–29) or the Periconiaceae (e.g., Periconia;Figs 42, 43),
but can be differentiated by their subglobose to hemispherical
ascomata with or without surrounding stromatic tissue, short
ascomatal necks without clypeus, cylindrical asci with a short
stipes, and narrowly fusiform, 1-septate, hyaline ascospores.
Summary of accepted families and genera
incertae sedis of the Massarineae
Based on our phylogenetic analyses and morphological evalu-
ation of fungi in the Massarineae, we recognise the following 12
families, as well as “Massarineae,incertae sedis”.
Bambusicolaceae. This family was established by Hyde
et al. (2013) to accommodate Bambusicola species on
bamboo (Dai et al. 2012). Palmiascoma on dead fronds of palms
E
D
B
A
C
Fig. 50. Monodictys capensis. A, B. Colonies on rice straw in culture; C. Co-
nidiophores and conidia; D, E. Conidia (E. bleached conidium). All from culture HR
1. Scale bars: A = 2 mm; B = 500 μm; C = 100 μm; D, E = 10 μm.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 127
in Thailand was also reported as a member of Bambusicolaceae
(Liu et al. 2015).
Dictyosporiaceae (Figs 2–5). Dictyosporiaceae (nom. prov.,
see Liu et al. 2015) includes the four asexual genera Aqua-
ticheirospora (Kodsueb et al. 2007), Dendryphiella (Jones et al.
2008), Dictyosporium (Fig. 2) and Pseudodictyosporium
(Kirschner et al. 2013), and two new genera with sexual morphs,
Gregarithecium (Fig. 3) and Pseudocoleophoma (Figs 4, 5). The
two asexual monotypic genera Cheirosporium (Cai et al. 2008)
and Kamatia (Kirschner et al. 2013) also belong in the Dictyo-
sporiaceae.“Diplococcium”asperum (Pirozynski 1972, Goh &
Hyde 1998) is located in the Dictyosporiaceae (Fig. 1), but the
type species of Diplococcium (D. spicatum) has phylogenetic
affinity with the Helotiales (Shenoy et al. 2010). This species as
well as Diplococcium pulneyense, the asexual morph of Otthia
pulneyensis (Subramanian & Sekar 1987), are both related to the
Dothideomycetes, and should be excluded from Diplococcium s.
str. Likewise, “Paraconiothyrium”flavescens in the Dictyospor-
iaceae clade should be renamed, because the type lineage of the
genus, represented by P. estuarinum, groups with the Didy-
mosphaeriaceae (= Montagnulaceae)(Verkley et al. 2014). The
Dictyosporiaceae clade includes Digitodesmium bambusicola
(Cai et al. 2002), but the phylogenetic placement of the generic
type D. elegans is unknown at present.
Didymosphaeriaceae (Figs 6–11). Ariyawansa et al. (2014)
accepted 16 genera including Karstenula (Fig. 6), Neokalmusia
(Figs 7, 8), and Paraphaeosphaeria (Fig. 9) as members of the
Didymosphaeriaceae, and subsequently three genera, Para-
camarosporium,Pseudocamarosporium, and Pseudotrichia were
also added to this family (Thambugala et al. 2014,
Wijayawardene et al. 2014d). We assigned the hyphomycetous
genus Spegazzinia (Figs 10, 11) with basauxic conidiogenesis
and pigmented conidia with spine-like appendages, to the Didy-
mosphaeriaceae, although it has long been treated as “Asco-
mycota,genera incertae sedis”(Wijayawardene et al. 2012).
However, taxonomic reassessment will be required for several
genera, such as Neokalmusia,Paraconiothyrium,Para-
phaeosphaeria and Pseudocamarosporium, because their
monophyletic status was not supported in this study. In addition to
these genera, the type species of Cucurbidothis (C. pityophila)
resides in the Didymosphaeriaceae (Fig. 1). Phaeodothis winteri
with a didymosphaeria-like sexual morph (Aptroot 1995), and
Sporidesmiella fusiformis with macronematous conidiophores
and obclavate, multi-distoseptate, pigmented conidia (Wu &
Zhuang 2005) are placed in the Didymosphaeriaceae, but se-
quences of the type species of these genera are presently un-
known. Two species of “Camarosporium”with muriform conidia,
C. brabeji and C. leucadendri (Marincowitz et al. 2008), are
positioned in the Didymosphaeriaceae, and have subsequently
been allocated to Pseudocamarosporium and Para-
camarosporium, respectively (Crous et al. 2015b). In contrast, the
type species of Camarosporium (C. quaternatum) is known to
cluster within the Pleosporineae (Crous et al. 2006,
Wijayawardene et al. 2014b). Munkovalsaria appendiculata
(Aptroot 2004) groups with Montagnula species (M. aloes,
M. opulenta and M. graminicola). However, Munkovalsaria is
apparently polyphyletic, because M. rubra is found outside the
Pleosporales (Voglmayr & Jaklitsch 2011, Hern
andez-Restrepo
et al. 2014). Although Munkovalsaria is regarded as a member
of the Dacampiaceae (Hyde et al. 2013), molecular data from the
generic type (M. donacina) are needed to clarify its phylogenetic
position.
Latoruaceae. This family was established by Crous et al.
(2015a) to accommodate two hyphomycetous genera, Latorua
and Polyschema, both mostly known from soil (Ellis 1976,
Shenoy et al. 2010, Crous et al. 2015a). More recently, a new
genus Matsushimamyces from soil has been added to this family
(Sharma et al. 2015).
Lentitheciaceae (Figs 12–24).Darksidea (Knapp et al.
2015), Katumotoa (Fig. 12), Keissleriella (Figs 13–18), Lenti-
thecium (Figs 19, 20), Murilentithecium (Wanasinghe et al.
2014), Phragmocamarosporium (Wijayawardene et al. 2015),
Poaceascoma (Phookamsak et al. 2015) and Tingoldiago
(Fig. 24) have been assigned in the Lentitheciaceae, and we add
another two genera, i.e., Neoophiosphaerella (Fig. 21) and
Setoseptoria (Figs 22, 23). The clade comprising species of
these six genera received moderate BP support (71 %; Fig. 1).
“Wettsteinina”lacustris is also a member of this clade (Schoch
et al. 2009). In our preliminary analysis using SSU and LSU
sequences, Ascorhombispora aquatica (Cai & Hyde 2007b)
resided in the Lentitheciaceae (data not shown), but the phylo-
genetic affinity of this species with members in the Dictyospor-
iaceae (Cai & Hyde 2007b)orDidymosphaeriaceae (Shearer
et al. 2009) has also been suggested. Further investigation us-
ing additional sequence data is needed to confirm familial
placement of this genus.
Macrodiplodiopsidaceae. Three species with coelomyce-
tous asexual morphs, i.e., Camarographium koreanum (on
Cornus,Verkley et al. 2005), Macrodiplodiopsis desmazieri (on
Platanus,Barr 1982), and Pseudochaetosphaeronema larense
(a human pathogen, Ahmed et al. 2014) comprise the Macro-
diplodiopsidaceae, which was recently established by Crous
et al. (2015a).
Massarinaceae (Figs 25–29).Byssothecium (Boise 1983),
Helminthosporium (Fig. 25), Massarina (Fig. 26), Stagonospora
(Figs 27–29) and Suttonomyces (Wijayawardene et al. 2015)
are accepted in the Massarinaceae (Fig. 1). Neottiosporina
paspali (Sutton & Alcorn 1974) should be treated as Stagono-
spora paspali (Atkinson 1897) based on the phylogenetic to-
pology (Fig. 1) as well as the morphological resemblance with
S. paludosa, the type species of Stagonospora (Quaedvlieg et al.
2013). The two Corynespora species C. leucadendri (Quaedvlieg
et al. 2013) and C. olivacea (Ellis 1960) clustered in this family
but are not congeneric. The type species of Corynespora
(C. mazei =C. cassiicola,Wei 1950) is sister of the clade of the
Pleosporineae +Massarineae (Hyde et al. 2013), hence the
Corynespora species in the Massarinaceae should be renamed.
“Didymosphaeria”spartii has been previously transferred to
Montagnula (Aptroot 1995) and also to Didymosphaerella
(Chlebicki 2009), but these generic placements are inappropriate
(Fig. 1). A cultural study of D. spartii (Scheinpflug 1958, using
CBS 183.58) indicated that it has a dendrophoma-like asexual
morph with conidia 4–7×1–2μm in size.
Morosphaeriaceae (Figs 30–39). In addition to Aquilomyces
(Fig. 30), Helicascus (Figs 35–37) and Morosphaeria (Figs 38,
39), which were previously recognised as members of the
Morosphaeriaceae (Suetrong et al. 2009, Hyde et al. 2013,
Knapp et al. 2015), we add the new genus Clypeoloculus
(Figs 31–34) to this family. Species in these four genera are only
known to have sexual morphs. A pleurophomopsis-like conidial
state has been reported for H. aquaticus (Zhang et al. 2013)as
its asexual morph, and we also observed this for H. aquaticus,as
well as for four species in Clypeoloculus, but these should be
regarded as spermatial morphs. “Pithomyces”valparadisiacus
TANAKA ET AL.
128
(Kirk 1983, Marincowitz et al. 2008) is basal to Clypeoloculus
species, and is found to be a rare case in which a species known
only from an asexual form belongs to the Morosphaeriaceae
(Fig. 1). A hyphomycetous genus Pithomyces is regarded as a
heterogenous group comprising more than three lineages
(Phookamsak et al. 2013), and the generic type (P. flavus) has
phylogenetic affinity with Astrosphaeriella basal to family Aigia-
laceae (Pratibha & Prabhugaonkar 2015).
Parabambusicolaceae (Figs 40, 41). This new family in-
cludes two new sexual genera, Aquastroma (Fig. 40) and Par-
abambusicola (Fig. 41). Multiseptospora formerly classified in
Pleosporales, genera incertae sedis (Liu et al. 2015) is also
accepted as a member of the Parabambusicolaceae. Two un-
named Monodictys species with muriform conidia are also
located in this clade, but this genus is obviously heterogenous
(see Notes in Monodictys capensis).
Periconiaceae (Figs 42, 43).Bambusistroma (Adam
cík et al.
2015), Flavomyces (Knapp et al. 2015), Periconia (Figs 42, 43)
and Noosia (Crous et al. 2011a) are accepted in this family.
“Sporidesmium”tengii is also placed in the Periconiaceae, but
the phylogenetic position of the type species of Sporidesmium
(S. atrum) is currently unknown. According to Shenoy et al.
(2006), this genus is not monophyletic and species in Spor-
idesmium are phylogenetically distributed in seven lineages
between two major ascomycete classes, Dothideomycetes and
Sordariomycetes.Periconia species used in this study did not
form a single clade. Therefore, a taxonomic revision of Periconia
species along with Noosia and “Sporidesmium”tengii should be
conducted in future.
Sulcatisporaceae (Figs 44–46).Magnicamarosporium
(Fig. 44)onDiplospora (Rubiaceae) and two species of
Sulcatispora,S. acerina (Fig. 45)onAcer (Aceraceae)and
S. berchemiae (Fig. 46)onBerchemia (Rhamnaceae), are
assigned to the Sulcatisporaceae.Neobambusicola (on Grewia sp.;
Malvaceae) previously placed in the Bambusicolaceae (Crous et al.
2014b) is accepted here as a member of the Sulcatisporaceae.
Trematosphaeriaceae (Fig. 47). This family was established
by Suetrong et al. (2011b) to accommodate the three genera,
Falciformispora,Halomassarina, and Trematosphaeria (Fig. 47)
(Hyde et al. 2013). Later, Bryosphaeria,Hadrospora, and Med-
icopsis were listed as additional members of the Trem-
atosphaeriaceae (Wijayawardene et al. 2014c). However, the
placement of Bryosphaeria has not been verified by molecular
data. Hadrospora may have affinity with the Lindgomycetaceae
(Tanaka, unpublished data), and Medicopsis belongs to the
Pleosporineae, rather than the Massarineae (Ahmed et al. 2014).
A coelomycetous species, Amarenographium solium with pig-
mented muriform conidia (Hodhod et al. 2012), has been re-
ported to have phylogenetic affinity to the Trematosphaeriaceae/
Didymosphaeriaceae, but molecular data from the generic type
(A. metableticum,Eriksson 1982) are currently unavailable.
Massarineae incertae sedis (Unknown Clades I–V; Figs
48–50).Fuscostagonospora (Fig. 48) on bamboo is located in
the Unknown Clade I and its familial placement remains unre-
solved. Pseudoxylomyces (Fig. 49), from freshwater environ-
ments, occupies a basal position in the Trematosphaeriaceae
(Unknown Clade II, Fig. 1). Bactrodesmium cubense (Unknown
Clade III), a sporodochial fungus with pigmented phragmoconidia
(Zucconi & Lunghini 1997), is a sister taxon of the Moro-
sphaeriaceae (Fig. 1). Monodictys capensis (Fig. 50), without a
known sexual morph (Mel’nik & Shabunin 2010), and Inflatispora
pseudostromatica, without a known asexual morph (Zhang et al.
2011), groups in Unknown Clade IV.
Morphological characteristics of sexual morphs
The Massarinaceae clade is supported by high BP support
(100 %, Fig. 1), but it is somewhat difficult to circumscribe based
on their sexual morphs. The Massarinaceae mainly contains two
types of sexual morphs; massarina-like and stagonospora-like.
The sexual morph observed in Helminthosporium (Fig. 25) and
Massarina (Fig. 26) have immersed, hemispherical, medium to
large (ca. 400–700 μm diam) ascomata with short papillate necks
covered by a clypeus, clavate asci, and broadly fusiform to
ellipsoidal, 1- to 3-septate, hyaline ascospores surrounded by a
thick conspicuous sheath. Those of Stagonospora (Figs 27–29)
are characterised by globose, small (ca. 200–300 μm diam)
ascomata lacking a clypeus, cylindrical to clavate asci, and
fusiform, 1-septate, hyaline ascospores with a thin sheath. This
family, however, includes further sexual species, such as “Didy-
mosphaerella (or Montagnula)”spartii with globose ascomata
and thick-walled, 1-septate, pigmented ascospores (Aptroot
1995, Chlebicki 2009), and Byssothecium circinans with sub-
globose ascomata with broadly papillate necks and versicoloured
ascospores (Boise 1983). Further molecular investigation using
additional taxa close to Massarina and Stagonospora may pro-
vide sufficient data to subdivide this family.
Bambusicolaceae,Macrodiplodiopsidaceae,Para-
bambusicolaceae and Sulcatisporaceae presently include only a
few species. One to two genera with sexual morphs are rec-
ognised in each family: Bambusicola (Dai et al. 2012) and Pal-
miascoma (Liu et al. 2015) in the Bambusicolaceae,
Macrodiplodiopsis (Crous et al. 2015a) in the Macro-
diplodiopsidaceae,Aquastroma (Fig. 40) and Parabambusicola
(Fig. 41) in the Parabambusicolaceae, and Sulcatispora (Figs 45,
46) in the Sulcatisporaceae. The sexual morphs in the Bambu-
sicolaceae,Parabambusicolaceae and Sulcatisporaceae are
similar to those of Massarina s. lat. (Aptroot 1998, Tanaka &
Harada 2003b) in having immersed to erumpent, hemispheri-
cal to depressed globose ascomata. Species in the Bambusi-
colaceae have conical ascomata with short ostiolar necks,
narrow pseudoparaphyses, cylindrical asci, and narrowly fusi-
form, 1-septate ascospores, whereas those in the Para-
bambusicolaceae lack a prominent neck, and have relatively
wide pseudoparaphyses, clavate to broadly cylindrical asci, and
clavate to fusiform ascospores with multiple transverse septa.
Species in the Sulcatisporaceae have subglobose ascomata,
trabeculate pseudoparaphyses, clavate asci, and broadly fusi-
form ascospores, and occur on woody host plant rather than
herbaceous host. On the other hand, the sexual morphs in the
Macrodiplodiopsidaceae are similar to those of Pleomassar-
iaceae s. lat.(Barr 1982). Macrodiplodiopsis desmazieri (=
Splanchnonema platani) in this family is characterised by large-
sized ascomata (500–900 μm diam) with thick ascomatal wall,
clavate asci, and dark brown ascospores with 3–5(–6) eudis-
tosepta (Barr 1982, Crous et al. 2015a).
Morosphaeriaceae (Figs 30–39) and Trematosphaeriaceae
(Fig. 47), mostly found from aquatic environments, are charac-
terised by immersed to erumpent, medium to large, black,
carbonaceous ascomata with or without papillate necks, but
necks are prominent and/or surrounded by a clypeus in mem-
bers of the Morosphaeriaceae. Asexual morphs are relatively
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 129
rare in these families with few exceptions, i.e., Pithomyces val-
paradisiacus (Morosphaeriaceae,Kirk 1983), and Trem-
atosphaeria grisea (Trematosphaeriaceae,Ahmed et al. 2014).
This tendency has been observed in the Aliquandostipitaceae
(Jahnulales), which also occurs in aquatic habitats, and a few
asexual species in the genera Brachiosphaera,Speiropsis, and
Xylomyces (Suetrong et al. 2011a).
We tentatively accept the Lentitheciaceae as a diverse
assemblage. Four groups seem to be recognised based on
morphological variations in the sexual morphs: Lentithecium with
immersed, globose ascomata without clypeus, and broadly
fusiform, 1-septate, hyaline ascospores (Figs 19, 20), Keissler-
iella with small, globose to subglobose ascomata with setose
necks and several septate ascospores (Figs 13–18), Seto-
septoria (Figs 22, 23) and Tingoldiago (Fig. 24) with single to
grouped, hemispherical ascomata and cylindrical to fusiform
ascospores, Katumotoa (Fig. 12) and Neoophiosphaerella
(Fig. 21) with subglobose ascomata without prominent necks and
fusiform or filiform ascospores. The morphological diversity of
the family, as well as relatively low phylogenetic support of the
clade (71 %, Fig. 1), strongly suggests polyphyly of the Lenti-
theciaceae s. lat. used in this study.
Similarly, various sexual morphs are observed in the Didy-
mosphaeriaceae (Figs 6–9). Most of the species in this family
have previously been placed in several genera of the Pleo-
sporineae, which include Didymosphaeria (Montagnula opulenta,
Phaeodothis winteri), Leptosphaeria (Paraconiothyrium fuckelii,
Neokalmusia scabrispora), and Phaeosphaeria (Neokalmusia
brevispora). The type species of Montagnula (M. infernalis;
Berlese 1896) has been treated as Leptosphaeria or Pleospora
(Wehmeyer 1961). Species having muriform ascospores similar
to Pleospora are known in Deniquelata (Ariyawansa et al. 2013)
and Tremateia (Kohlmeyer et al. 1995). Sexual morphs in the
Didymosphaeriaceae have small to medium, globose to sub-
globose ascomata with papillate necks and melanised, didymo/
phragmo/dictyo-ascospores resembling those of the Pleospor-
ineae, but the ascomata are surrounded by abundant hyphae
that form an apical clypeus at times, as was defined by Barr
(2001). In addition, Bimuria (Hawksworth et al. 1979), a soil-
borne ascomycete, with unique morphological characters such
as ascomata composed of very thin peridium, 2-spored asci, and
muriform, dark brown, verrucose ascospores, also belongs in the
Didymosphaeriaceae. The family includes a wide variety of
sexual morphs with dark coloured ascospores, as well as
coniothyrium-like or Spegazzinia asexual morphs, and the clade
receives 99 % BP support (Fig. 1).
In contrast, the Dictyosporiaceae and Periconiaceae mostly
comprise asexual species. The few known sexual morphs in
these families are characterised by single to grouped, globose to
hemispherical ascomata with or without surrounding stromatic
tissue, cylindrical asci with short stipes, and fusiform to broadly
fusiform, 1-septate, hyaline ascospores. Examples are Dictyo-
sporium (e.g., D. meiosporum;Liu et al. 2015), Gregarithecium
(Fig. 3) and Pseudocoleophoma (Figs 4, 5) in the Dictyospor-
iaceae and Bambusistroma (Adam
cík et al. 2015) and Periconia
(Figs 42, 43) in the Periconiaceae. These morphological features
agree in most respects with those of Massarina s. lat. (Aptroot
1998, Tanaka & Harada 2003b) and are similar to those of
species in the Massarinaceae (e.g., Stagonospora). In the
Latoruaceae, no sexual morphs are known at present (Crous
et al. 2015a).
Morphological characteristics of asexual
morphs
Although various asexual genera are found in the Massarineae,
hyphomycetous species with macronematous, mononematous
conidiophores are restricted to the Massarinaceae [Helmintho-
sporium (Fig. 25) and Corynespora], Periconiaceae [Periconia
(Fig. 43) and Sporidesmium], Dictyosporiaceae (Diplococcium
and Dendryphiella), and Didymosphaeriaceae (Sporidesmiella).
Similarly, synnematous hyphomycetes are extremely rare in this
suborder and have only been observed in one instance, in
Aquaticheirospora (Kodsueb et al. 2007) in the Dictyospor-
iaceae. Hyphomycetous asexual morphs without prominent co-
nidiophores are found in the Periconiaceae (Noosia),
Latoruaceae (Latorua and Polyschema), and the Moro-
sphaeriaceae (Pithomyces). Sporodochial hyphomycetes are
scattered through the Dictyosporiaceae [Dictyosporium (Fig. 2)
and its morphologically similar genera], Didymosphaeriaceae
(Spegazzinia,Figs 10, 11), Parabambusicolaceae (Monodictys),
and Unknown Clades III (Bactrodesmium) and IV (Monodictys,
Fig. 50). In contrast, coelomycetous genera seem to be more
common than hyphomycetous asexual morphs and are present
throughout this suborder, i.e., in the Bambusicolaceae (Bambu-
sicola), Dictyosporiaceae (Pseudocoleophoma,Figs 4, 5),
Didymosphaeriaceae [Paracamarosporium,Paraconiothyrium,
Pseudocamarosporium,Karstenula,Paraphaeosphaeria
(Fig. 9)], Lentitheciaceae [Keissleriella (Figs 13, 16), Seto-
septoria], Macrodiplodiopsidaceae (Camarographium,Macro-
diplodiopsis,Pseudochaetosphaeronema), Massarinaceae
(Stagonospora,Figs 27–29), Sulcatisporaceae [Magnicamar-
osporium (Fig. 44), Sulcatispora (Figs 45, 46)] and Unknown
Clades I (Fuscostagonospora,Fig. 48).
In general, Dictyosporiaceae is characterised by hyphomy-
cetous asexual morphs with pigmented cheiroconidia. These
usually have sporodochial (e.g., Dictyosporium,Digitodesmium,
and Pseudodictyosporium) or synnematous conidiomata (e.g.,
Aquaticheirospora). Dendryphiella and Diplococcium have
macronematous, mononematous conidiophores with pigmented,
multi-celled, tretic conidia. The coelomycetous morphs known in
this family are “Paraconiothyrium”flavescens and Pseudoco-
leophoma species, with phialidic, 1-celled conidia. In contrast,
most asexual species in the Periconiaceae have mono-
nematous, macronematous conidiophores and produce conidia
in chains (e.g., Periconia), but some deviating species, such as
“Sporidesmium”tengii have monoblastic, obpyriform, multi-
septate, pigmented conidia (Wu & Zhuang 2005), and Noosia
which lacks a prominent conidiophore (Crous et al. 2011a), are
also known in the family. Further phylogenetic investigations of
asexual genera exhibiting morphological resemblance to these
two groups but without molecular information, such as
dictyosporium-like (e.g., Cheiromycella,Digitomyces,Para-
tetraploa;Seifert et al. 2011) and periconia-like genera (e.g.,
Lacellinopsis,Sadasivania,Trichobotrys;Seifert et al. 2011),
should be conducted to evaluate the phylogenetic significance
of these phenotypic characters at familial level, and/or to clarify
taxonomic understanding of the Dictyosporiaceae and Peri-
coniaceae. Although in some families (e.g., the Didymos-
phaeriaceae and Massarinaceae), asexual morphs are
extremely diverse and thus asexual characters alone appear
insufficient for familial circumscriptions, there are many recent
examples of families that are well defined by asexual
TANAKA ET AL.
130
morphological characters along with their phylogenetic infor-
mation. These include the Coniothyriaceae (De Gruyter et al.
2013), Cladosporiaceae (Bensch et al. 2012), Kirsch-
steiniotheliaceae (Boonmee et al. 2012), Planistromellaceae
(Minnis et al. 2012), and Tetraplosphaeriaceae (Tanaka et al.
2009).
Habitat
In contrast to the Pleosporineae which comprises numerous
plant pathogens (Zhang et al. 2009b), such as necrotrophs (e.g.,
Alternaria,Bipolaris) and hemibiotrophs (e.g., Leptosphaeria,
Setosphaeria) on economically important crops (Ohm et al.
2012), the Massarineae, a sister lineage of the Pleosporineae,
mostly includes saprobes on various plant substrates.
Species in the Morosphaeriaceae and Trematosphaeriaceae
have been reported mostly as saprobes on decomposed woody
substrates submerged in freshwater or marine habitats.
Recently, several human pathogenic species have been reported
in genera in the Trematosphaeriaceae, and an association of the
virulence factors with oligotrophism or halotolerance has been
suggested (Ahmed et al. 2014). This may further indicate the
presence of undescribed lineages of mycetoma agents in the
Morosphaeriaceae.
Most other families in the Massarineae do not seem to have
specific habitat preferences in each family. Species in the Dic-
tyosporiaceae, for example, are reported from ecologically
diverse environments, i.e., terrestrial monocots and dicots (e.g.,
Dictyosporium strelitziae and Diplococcium asperum;Pirozynski
1972, Crous et al. 2009a), submerged wood in freshwater (e.g.,
Aquaticheirospora lignicola;Kodsueb et al. 2007), mangroves or
drift wood in marine ecosystems (Dictyosporium inflatum;
Kirschner et al. 2013), and rhizosphere soil (Pseudodictyospo-
rium elegans;De Gruyter et al. 2013). In the Didymosphaer-
iaceae, many coelomycetous species similar to Phoma s. lat. are
generally soil-borne fungi (Verkley et al. 2014), but species with
other ecological features, e.g., endophytes (Dendrothyrium;
Verkley et al. 2014), mycoparasites (Paraphaeosphaeria mini-
tans;Campbell 1947), symbionts with scale insects (Cucurbi-
dothis pityophila;Casagrande 1969, Barr 1990b), and marine
saprobes (Tremateia halophila;Kohlmeyer et al. 1995) are also
known. Paraconiothyrium fuckelii in the Didymosphaeriaceae is
known as an agent of mycetoma and a serious plant pathogen of
the Rosaceae (Verkley et al. 2014).
In the suborder Massarineae, relatively few species are
known as plant pathogens. They are restricted to the Didy-
mosphaeriaceae (Paraconiothyrium tiliae,Deniquelata barring-
toniae;Butin & Kehr 1995, Ariyawansa et al. 2013), Massarineae
(Byssothecium circinans,Stagonospora spp., Helminthosporium
solani;Kaiser et al. 1979, Semeniuk 1983, Errampalli et al. 2001)
and Periconiaceae (Periconia igniaria;Kolomiets et al. 2008);
lineages rich in asexual species. Coprophilous species such as
those in the Sporormiaceae (Kruys & Wedin 2009) and lichen-
icolous species such as Arthopyrenia salicis (Nelsen et al. 2009)
are not known from the Massarineae.
Future studies
We examined the morphology of 106 taxa belonging to the
Massarineae and analysed their phylogenetic relationships
based on sequences from SSU rDNA, LSU rDNA and tef1, along
with sequences of 131 taxa previously assigned to the Mas-
sarineae. Our results delineated 10 new genera and 29 new
species/new combinations in more than 12 families and five
unknown lineages (Fig. 1). Our study has contributed to the
understanding of species diversity within the Massarineae, and
improves the classification of these species, but several taxo-
nomic issues remain unclear. The Lentitheciaceae s. lat. used
here, for example, is probably polyphyletic based on the
ecological and morphological divergence of these species, as
well as based on our phylogenetic analysis, although we have
tentatively accepted the family at this point. Furthermore, the
presence of several lineages (Unknown Clades I to V), which we
were unable to assign to any existing families, makes it certain
that there are many undiscovered taxa, which should form a new
family in the suborder.
Several comprehensive works have been published recently
that have shown interest in the phylogeny of the Dothideomy-
cetes.Schoch et al. (2006) provided a fundamental overview of
the class, and its composition of two subclasses, the Pleo-
sporomycetidae (pseudoparaphyses present) and the Dothi-
deomycetidae (pseudoparaphyses absent), based on
phylogenetic analyses of four loci (nucSSU, nucLSU rDNA,
tef1,rpb2) from 96 taxa. Schoch et al. (2009) reconstructed the
dothideomycete phylogeny based on five genes (nucSSU,
nucLSU rDNA, tef1,rpb1,rpb2) from 356 isolates, and dis-
cussed the evolutionary transitions of ecological characteristics.
Following these publications, the understanding of the natural
relationships among dothideomycetous taxa and their taxo-
nomic revision have tremendously progressed, and the number
of accepted families in this class has been increased from 41
(Schoch et al. 2009)to105(Hyde et al. 2013)withorwithout
molecular evidence. Monographic revision based on the type
specimens along with phylogeneric analyses have been pub-
lished by Zhang et al. (2012) who concentrated on reassess-
ment of genera in the Pleosporales, and by Hyde et al. (2013)
who circumscribed each family in the Dothideomycetes.The
phylogenies of plant pathogenic taxa in this class, such as
Alternaria (Woudenberg et al. 2013), Cercospora (Groenewald
et al. 2013), Cladosporium (Bensch et al. 2012), Phoma (De
Gruyter et al. 2013), Pseudocercospora (Crous et al. 2013a),
Septoria (Quaedvlieg et al. 2013, Verkley et al.2013)and
several genera in the Botryosphaeriales (Phillips et al. 2013,
Slippers et al. 2013, Wikee et al.2013), have been inten-
sively studied. More recently, a special issue of Phytotaxa has
been published, comprising 26 articles focused on the taxon-
omy and phylogeny of the Dothideomycetes (Mckenzie et al.
2014).
However, these works on Dothideomycetes include relatively
few species belonging to the Massarineae. The number of
Massarineae taxa used in phylogenetic analyses of the Dothi-
deomycetes was only six (among 96 taxa; Schoch et al. 2006),
and 35 (among 356 taxa; Schoch et al. 2009). In the revision of
the Pleosporales by Zhang et al. (2012) who recognised the
validity of Massarineae, only 46 taxa of the suborder were
phylogenetically analysed. Furthermore, most families in this
suborder including the Bambusicolaceae (Hyde et al. 2013),
Lentitheciaceae (Zhang et al. 2009b), Morosphaeriaceae
(Suetrong et al. 2009), and Trematosphaeriaceae (Suetrong
et al. 2011b), have been lineages recently recognised based
on molecular data, and are currently characterised morphologi-
cally by only a few genera. To provide more precise circum-
scription of these families, as well as of the unnamed lineages we
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 131
have found (Clades I to V, Fig. 1), further phylogenetic analyses
using a greater number of taxa should be conducted with priority.
These should include a fundamental taxonomic approach to
species discovery along with sequence verification, such as the
work undertaken by Fungal Planet (Crous et al. 2015c), Fungal
Diversity Notes (Liu et al. 2015) and Fungal Systematics and
Evolution (FUSE) (Crous et al. 2015b).
In due course further phylogenetic study is also necessary of
the more than 1 500 asexual genera treated as “Ascomycota,
genera incertae sedis”(Wijayawardene et al. 2012); those
without sexual links or molecular phylogenetic information.
However, asexual characters alone may not provide good res-
olution for familial circumscription for some aquatic lineages,
such as Morosphaeriaceae and Trematosphaeriaceae, in which
predominantly sexual species are known. In contrast, species in
the Dictyosporiaceae and Periconiaceae comprise a high pro-
portion of asexual taxa. These have previously been treated as
Massarinaceae (Hyde et al. 2013), but we have further charac-
terised these as independent families based on their sexual
morphs, in addition to their distinct asexual morphologies.
Recognition of the Dictyosporiaceae and Periconiaceae partially
indicates the phylogenetic significance of asexual taxa, but
further comprehensive taxonomic work based on the holomorph
and not weighted towards a particular fungal morph (e.g., Crous
et al. 2009b, Tanaka et al. 2009, 2010, Voglmayr & Jaklitsch
2011, Dai et al. 2012, Phillips et al. 2013, Boonmee et al.
2014) should be conducted to revise sexual morph-based
fungal systematics.
ACKNOWLEDGEMENTS
We gratefully acknowledge N. Asama, K. Izumi, T. Handa, S. Hatakeyama, Y.
Hiro, N. Nakagawara, A. Ooba, Y. Ooki, J. Onodera, and Y. Suzuki for providing
fungal specimens. We thank the curator of TMI, E. Nagasawa, who permitted us
to examine type collections. We wish to acknowledge V.A. Mel'nik for kindly
supplying the specimen of Monodictys capensis and H.A. Raja for kindly
providing literature of Periconiaceae. We also thank editors and anonymous
reviewers for critically reviewing the manuscript. This work was partially sup-
ported by grants from the Japan Society for the Promotion of Science (JSPS
25440199 and 26291084) and Hirosaki University Grant for Exploratory Research
by Young Scientists and Newly-appointed Scientists for financial support
(2010–2013, 2015).
REFERENCES
Adam
cík S, Cai L, Chakraborty D, et al. (2015). Fungal biodiversity profiles
1–10. Cryptogamie Mycologie 36: 121–166.
Ahmed SA, van de Sande WWJ, Stevens DA, et al. (2014). Revision of agents
of black-grain eumycetoma in the order Pleosporales.Persoonia 33:
141–154.
Akaike H (1974). A new look at the statistical model identification. IEEE
Transactions on Automatic Control 19: 716–723.
Anastasiou CJ (1963). Fungi from salt lakes. II. Ascomycetes and fungi
imperfecti from the Salton sea. Nova Hedwigia 6: 243–276.
Aptroot A (1995). Redisposition of some species excluded from Didymosphaeria
(Ascomycotina). Nova Hedwigia 60: 325–379.
Aptroot A (1998). A world revision of Massarina (Ascomycota). Nova Hedwigia
66:89–162.
Aptroot A (2004). Two new ascomycetes with long gelatinous appendages
collected from monocots in the tropics. Studies in Mycology 50:
307–311.
Ariyawansa HA, Maharachchikumbura SSN, Karunarathne SC, et al. (2013).
Deniquelata barringtoniae gen. et sp. nov., associated with leaf spots of
Barringtonia asiatica.Phytotaxa 105:11–20.
Ariyawansa HA, Tanaka K, Thambugala KM, et al. (2014). A molecular phylo-
genetic reappraisal of the Didymosphaeriaceae (= Montagnulaceae). Fungal
Diversity 68:69–104.
Atkinson GF (1897). Some fungi from Alabama collected chiefly during the years
1889–1892. Bulletin of the Cornell University (Science) 3:1–50.
Barbosa FR, Gusm~
ao LFP (2011). Conidial fungi from semi-arid Caatinga Biome
of Brazil. Rare freshwater hyphomycetes and other new records. Myco-
sphere 2: 475–485.
Barr ME (1979). A classification of Loculoascomycetes.Mycologia 71: 935–957.
Barr ME (1982). On the Pleomassariaceae (Pleosporales) in North America.
Mycotaxon 15: 349–383.
Barr ME (1987). Prodromus to class Loculoascomycetes. Published by the
author, Amherst, Massachusetts.
Barr ME (1990a). Melanommatales (Loculoascomycetes). North American Flora,
Series II 13:1–129.
Barr ME (1990b). Some dictyosporous genera and species of Pleosporales in
North America. Memoirs of the New York Botanical Garden 62:1–92.
Barr ME (1992). Notes on the Lophiostomataceae (Pleosporales). Mycotaxon
45: 191–221.
Barr ME (2001). Montagnulaceae, a new family in the Pleosporales, and lec-
totypification of Didymosphaerella.Mycotaxon 77: 193–200.
Bensch K, Braun U, Groenewald JZ, et al. (2012). The genus Cladosporium.
Studies in Mycology 72:1–401.
Berlese AN (1894). Icones fungorum omnium hucusque cognitorum, vol. 1
(Reprint Bibliotheca Mycologica 16A:1–243).
Berlese AN (1896). Icones fungorum omnium hucusque cognitorum, vol. 2
(Reprint Bibliotheca Mycologica 16B:1–216).
Boise J (1983). On Trematosphaeria circinans and reinstatement of the genus
Byssothecium.Mycologia 75: 666–669.
Boise J (1985). New combinations in the Pleomassariaceae and the Massar-
inaceae.Mycotaxon 22: 477–482.
Boonmee S, KoKo TW, Chukeatirote E, et al. (2012). Two new Kirsch-
steiniothelia species with Dendryphiopsis anamorphs cluster in Kirsch-
steiniotheliaceae fam. nov. Mycologia 104: 698–714.
Boonmee S, Rossman AY, Liu JK, et al. (2014). Tubeufiales,ord.nov.,
integrating sexual and asexual generic names. Fungal Diversity 68:
239–298.
Booth C (1968). Didymosphaeria igniaria sp. nov., the perfect state of Periconia
igniaria.Transactions of the British Mycological Society 51: 803–805.
Bose SK (1961). Studies on Massarina Sacc. and related genera. Phytopa-
thologische Zeitschrift 41: 151–213.
Bunning SE, Griffiths DA (1984). Spore development in species of Periconia II.
P. byssoides and P. igniaria.Transactions of the British Mycological Society
82: 397–404.
Butin H, Kehr R (1995). Leaf blotch of lime associated with Asteromella tiliae
comb. nov. and the latter's connection to Didymosphaeria petrakiana.
Mycological Research 99: 1191–1194.
Cai L, Guo XY, Hyde KD (2008). Morphological and molecular characterisation
of a new anamorphic genus Cheirosporium, from freshwater in China.
Persoonia 20:53–58.
Cai L, Hyde KD (2007a). Anamorphic fungi from freshwater habitats in China:
Dictyosporium tetrasporum and Exserticlava yunnanensis spp. nov., and two
new records for Pseudofuscophialis lignicola and Pseudobotrytis terrestris.
Mycoscience 48: 290–296.
Cai L, Hyde KD (2007b). Ascorhombispora aquatica gen. et sp. nov. from a
freshwater habitat in China, and its phylogenetic placement based on mo-
lecular data. Cryptogamie Mycologie 28: 291–300.
Cai L, Zhang K, McKenzie EHC, et al. (2002). Acrodictys liputii sp. nov. and
Digitodesmium bambusicola sp. nov. from bamboo submerged in the Liput
River in the Philippines. Nova Hedwigia 75: 525–532.
Cai L, Zhang K, McKenzie EHC, et al. (2003). New species of Canalisporium
and Dictyosporium from China and a note on the differences between these
genera. Cryptogamie Mycologie 24:3–11.
C^
amara MPS, O'Neill NR, van Berkum P, et al. (2000). Ophiosphaerella agrostis
sp. nov. and its relationship to other species of Ophiosphaerella.Mycologia
92: 317–325.
C^
amara MPS, Palm ME, van Berkum P, et al. (2001). Systematics of Para-
phaeosphaeria: a molecular and morphological approach. Mycological
Research 105:41–56.
C^
amara MPS, Palm ME, van Berkum P, et al. (2002). Molecular phylogeny of
Leptosphaeria and Phaeosphaeria.Mycologia 94: 630–640.
C^
amara MPS, Ramaley AW, Castlebury LA, et al. (2003). Neophaeosphaeria
and Phaeosphaeriopsis, segregates of Paraphaeosphaeria.Mycological
Research 107: 516–522.
TANAKA ET AL.
132
Campbell J, Ferrer A, Raja HA, et al. (2007). Phylogenetic relationships among
taxa in the Jahnulales inferred from 18S and 28S nuclear ribosomal DNA
sequences. Canadian Journal of Botany 85: 873–882.
Campbell J, Shearer CA, Mitchell JI, et al. (2002). Corollospora revisited: a mo-
lecular approach. In: (Hyde KD, ed), Fungi in marine environments. Fungal
diversity research series 7Fungal Diversity Press, Hong Kong: 15–33.
Campbell WA (1947). A new species of Coniothyrium parasitic on sclerotia.
Mycologia 39: 190–195.
Casagrande F (1969). Ricerche biologiche e sistematische su particolari
ascomiceti pseudosferiali. Phytopathologische Zeitschrift 66:97–136.
Chen JL, Hwang CH, Tzean SS (1991). Dictyosporium digitatum, a new
hyphomycete from Taiwan. Mycological Research 95: 1145 –1149.
Chlebicki A (2009). Fungi on higher plants of the upper limit of the alpine zone:
new species from Tian Shan. Mycotaxon 110: 443–450.
Clements FE, Shear CL (1931). The genera of fungi. Wilson, New York.
Constantinescu O (1993). Teleomorph-anamorph connections in ascomycetes:
Microdiplodia anamorph of Karstenula rhodostoma.Mycological Research
97: 377–380.
Corbaz R (1956). Recherches sur le genre Didymella Sacc. Phytopathologische
Zeitschrift 28: 375–414.
Corlett M, Smith JD (1978). Didymella proximella and its Stagonospora ana-
morph. Canadian Journal of Botany 56: 2818–2824.
Crane JL, Shearer CA (1991). A nomenclator of Leptosphaeria V. Cesati & G. de
Notaris (Mycota-Ascomycotina-Loculoascomycetes). Illinois Natural History
Survey Bulletin 34:1–355.
Crous PW, Braun U, Hunter GC, et al. (2013a). Phylogenetic lineages in
Pseudocercospora.Studies in Mycology 75:37–114.
Crous PW, Braun U, Wingfield MJ, et al. (2009a). Phylogeny and taxonomy of
obscure genera of microfungi. Persoonia 22: 139–161.
Crous PW, Carris LM, Giraldo A, et al. (2015a). The Genera of Fungi –fixing
the application of the type species of generic names –G2:Allantopho-
mopsis,Latorua,Macrodiplodiopsis,Macrohilum,Milospium,Protostegia,
Pyricularia,Robillarda,Rotula,Septoriella,Torula, and Wojnowicia.IMA
Fungus 6: 163–198.
Crous PW, Gams W, Stalpers JA, et al. (2004). MycoBank: an online initiative to
launch mycology into the 21st century. Studies in Mycology 50:19–22.
Crous PW, Groenewald JZ (2013). A phylogenetic re-evaluation of Arthrinium.
IMA Fungus 4: 133–154.
Crous PW, Groenewald JZ, Lombard L, et al. (2012). Homortomyces gen. nov.,
a new dothidealean pycnidial fungus from the Cradle of Humankind. IMA
Fungus 3: 109–115.
Crous PW, Groenewald JZ, Shivas RG, et al. (2011a). Fungal Planet description
sheets: 69–91. Persoonia 26: 108 –156.
Crous PW, Schoch CL, Hyde KD, et al. (2009b). Phylogenetic lineages in the
Capnodiales.Studies in Mycology 64:17–47.
Crous PW, Schumacher RK, Wingfield MJ, et al. (2015b). Fungal Systematics
and Evolution: FUSE 1. Sydowia 67:81–118.
Crous PW, Shivas RG, Quaedvlieg W, et al. (2014a). Fungal Planet description
sheets: 214–280. Persoonia 32: 184 –306.
Crous PW, Slippers B, Wingfield MJ, et al. (2006). Phylogenetic lineages in the
Botryosphaeriaceae.Studies in Mycology 55: 235–253.
Crous PW, Summerell BA, Carnegie AJ, et al. (2007). Foliicolous Mycos-
phaerella spp. and their anamorphs on Corymbia and Eucalyptus.Fungal
Diversity 26: 143–185.
Crous PW, Summerell BA, Shivas RG, et al. (2011b). Fungal Planet description
sheets: 92–106. Persoonia 27: 130 –162.
Crous PW, Wingfield MJ, Guarro J, et al. (2013b). Fungal Planet description
sheets: 154–213. Persoonia 31: 188 –296.
Crous PW, Wingfield MJ, Guarro J, et al. (2015c). Fungal Planet description
sheets: 320–370. Persoonia 34: 167 –266.
Crous PW, Wingfield MJ, Schumacher RK, et al. (2014b). Fungal Planet
description sheets: 281–319. Persoonia 33: 212 –289.
Dai DQ, Bhat DJ, Liu JK, et al. (2012). Bambusicola, a new genus from bamboo
with asexual and sexual morphs. Cryptogamie Mycologie 33: 363–379.
Day MJ, Gibas CFC, Fujimura KE, et al. (2006). Monodictys arctica, a new
hyphomycete from the roots of Saxifraga oppositifolia collected in the Ca-
nadian High Arctic. Mycotaxon 98: 261–272.
De Gruyter J, Aveskamp MM, Woudenberg JHC, et al. (2009). Molecular
phylogeny of Phoma and allied anamorph genera: towards a reclassification
of the Phoma complex. Mycological Research 113: 508 –519.
De Gruyter J, Woudenberg JHC, Aveskamp MM, et al. (2013). Redisposition of
Phoma-like anamorphs in Pleosporales.Studies in Mycology 75:1–36.
Dennis RWG (1978). British Ascomycetes,3
rd
edn. J. Cramer, Vaduz.
Ellis MB (1960). Dematiaceous hyphomycetes. I. Mycological Papers 76:1–36.
Ellis MB (1961). Dematiaceous hyphomycetes. III. Mycological Papers 82:1–55.
Ellis MB (1971). Dematiaceous hyphomycetes. Commonwealth Mycological
Institute, Kew, Surrey, UK.
Ellis MB (1976). More dematiaceous hyphomycetes. Commonwealth Mycolog-
ical Institute, Kew, Surrey, UK.
Eriksson OE (1967). On graminicolous pyrenomycetes from Fennoscandia. 2.
Phragmosporous and scolecosporous species. Arkiv för Botanik 6:
381–440.
Eriksson OE (1981). The families of bitunicate ascomycetes. Opera Botanica 60:
1–220.
Eriksson OE (1982). Notes on ascomycetes and coelomycetes from NW
Europe. Mycotaxon 15: 189–202.
Eriksson OE (1989). NaClO, sodium hypochlorite, a powerful agent in studies of
ascospore morphology. Systema Ascomycetum 8:29–57.
Eriksson OE, Hawksworth DL (2003). Saccharicola, a new genus for two
Leptosphaeria species on sugar cane. Mycologia 95: 426–433.
Eriksson OE, Winka K (1998). Families and higher taxa of Ascomycota.Myconet
1:17–24.
Errampalli D, Saunders JM, Holley JD (2001). Emergence of silver scurf (Hel-
minthosporium solani) as an economically important disease of potato. Plant
Pathology 50: 141–153.
Goh TK, Ho WH, Hyde KD, et al. (1997). Four new species of Xylomyces from
submerged wood. Mycological Research 101: 1323–1328.
Goh TK, Hyde KD (1998). A synopsis of and a key to Diplococcium species,
based on the literature, with a description of a new species. Fungal Diversity
1:65–83.
Goh TK, Hyde KD, Ho WH, et al. (1999). A revision of the genus Dictyosporium,
with descriptions of three new species. Fungal Diversity 2:65–100.
Goos RD, Brooks RD, Lamore BJ (1977). An undescribed hyphomycete from
wood submerged in a Rhode Island stream. Mycologia 69: 280–286.
Grandaubert J, Lowe RGT, Soyer JL, et al. (2014). Transposable element-
assisted evolution and adaptation to host plant within the Leptosphaeria
maculans-Leptosphaeria biglobosa species complex of fungal pathogens.
BMC Genomics 15: 891.
Groenewald JZ, Nakashima C, Nishikawa J, et al. (2013). Species concepts in
Cercospora: spotting the weeds among the roses. Studies in Mycology 75:
115 –170.
Grove WB (1935). British stem and leaf-fungi (Coelomycetes), Vol. I. Sphaer-
opsidales. Cambridge University Press, Cambridge, UK.
Han JG, Hosoya T, Sung GH, et al. (2014). Phylogenetic reassessment of
Hyaloscyphaceae sensu lato (Helotiales,Leotiomycetes) based on multi-
gene analyses. Fungal Biology 118: 150–167.
Hawksworth DL, Kirk PM, Sutton BC, et al. (1995). Ainsworth and Bisby's
Dictionary of the Fungi,8
th
edn. CAB International, Wallingford, UK.
Hawksworth DL, Sutton BC, Ainsworth GC (1983). Ainsworth and Bisby's Dic-
tionary of the Fungi,7
th
edn. Commonwealth Mycological Institute, Kew, UK.
Hawksworth DL, Yen CC, Sheridan JE (1979). Bimuria novae-zelandiae gen. et
sp. nov., a remarkable ascomycete isolated from a New Zealand barley field.
New Zealand Journal of Botany 17: 267–273.
Hedjaroude GA (1968). Etudes taxonomiques sur les Phaeosphaeria Miyake et
leurs formes voisines (Ascomycetes). Sydowia 22:57–107.
Hern
andez-Restrepo M, Casta~
neda-Ruiz RF, Gen
eJ,et al. (2014). Two new
species of Solicorynespora from Spain. Mycological Progress 13:
157–164.
Hirayama K, Tanaka K (2011a). Notes on two species of Massarina sensu lato
from Island Iriomote in Japan (in Japanese). Aomorisizenshikenkyuu 16:
43–47.
Hirayama K, Tanaka K (2011b). Taxonomic revision of Lophiostoma and
Lophiotrema based on reevaluation of morphological characters and mo-
lecular analyses. Mycoscience 52: 401–412.
Hirayama K, Tanaka K, Raja HA, et al. (2010). A molecular phylogenetic
assessment of Massarina ingoldiana sensu lato.Mycologia 102: 729–746.
Hirayama K, Yonezawa H, Tanaka K (2012). Description of Hyphomycetes,
Dictyosporium bulbosum and D. digitatum, new to Japan (in Japanese).
Aomorisizensikenkyu 17:51–54.
Hodhod MS, Abdel-Wahab MA, Bahkali AHA, et al. (2012). Amarenographium
solium sp. nov. from Yanbu mangroves in the Kingdom of Saudi Arabia.
Cryptogamie Mycologie 33: 285–294.
Holm L (1957).
Etudes taxonomiques sur les pl
eosporac
ees. Symbolae Bota-
nicae Upsalienses 14:1–188.
Hosoya T, Huhtinen S (2002). Hyaloscyphaceae in Japan (7): Hyaloscypha
albohyalina var. monodictys var. nov. Mycoscience 43: 405 –409.
Hsieh WH (1979). The causal organism of sugarcane leaf blight. Mycologia 71:
892–898.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 133
Hu DM, Cai L, Chen H, et al. (2010). Four new freshwater fungi associated with
submerged wood from Southwest Asia. Sydowia 62: 191–203.
Hughes SJ (1953a). Conidiophores, conidia, and classification. Canadian
Journal of Botany 31: 577–659.
Hughes SJ (1953b). Fungi from the Gold Coast. II. Mycological Papers 50:
1–104.
Huhndorf SM (1992). Studies in Leptosphaeria. Transfer of Leptosphaeria
opuntiae to Montagnula (Ascomycetes). Brittonia 44: 208–212.
Hyde KD (1991). Massarina velatospora and a new mangrove-inhabiting spe-
cies, M. ramunculicola sp. nov. Mycologia 83: 839–845.
Hyde KD (1995). The genus Massarina, with a description of M. eburnea and an
annotated list of Massarina names. Mycological Research 99: 291–296.
Hyde KD, Aptroot A (1998). Tropical freshwater species of the genera Massarina
and Lophiostoma (ascomycetes). Nova Hedwigia 66: 489–502.
Hyde KD, Fröhlich J, Taylor JE (1998). Fungi from palms. XXXVI. Reflections on
unitunicate ascomycetes with apiospores. Sydowia 50:21–80.
Hyde KD, Goh TK (1999). Tropical Australian freshwater fungi. XVI. Some new
melanommataceous fungi from woody substrata and a key to genera of
lignicolous loculoascomycetes in freshwater. Nova Hedwigia 68: 251–272.
Hyde KD, Jones EBG, Liu JK, et al. (2013). Families of Dothideomycetes.
Fungal Diversity 63:1–313.
Hyde KD, McKenzie EHC, KoKo TW (2011). Towards incorporating anamorphic
fungi in a natural classification –checklist and notes for 2010. Mycosphere
2:1–88.
Hyde KD, Wong WSW, Aptroot A (2002). Marine and estuarine species of
Lophiostoma and Massarina. In: (Hyde KD, ed), Fungi in marine environ-
ments. Fungal diversity research series 7Fungal Diversity Press, Hong
Kong: 93–109.
Jobb G (2011). Treefinder Mar 2011. Available at: http://www.treefinder.de.
Jones EBG, Klaysuban A, Pang KL (2008). Ribosomal DNA phylogeny of
marine anamorphic fungi: Cumulospora varia,Dendryphiella species and
Orbimyces spectabilis.The Raffles Bulletin of Zoology supplement 19:
11–18.
Kaiser WJ, Ndimande BN, Hawksworth DL (1979). Leaf-scorch disease of
sugarcane in Kenya caused by a new species of Leptosphaeria.Mycologia
71: 479–492.
Karsten PA (1873). Mycologia Fennica. Pars secunda. Pyrenomycetes. Bidrag
till K€
annedom af Finlands Natur och Folk 23:1–252.
Khashnobish A, Shearer CA (1993). Reexamination of eight Leptosphaeria
species (Ascomycetes:Pleosporales) originally described on Typha.Nova
Hedwigia 57:33–46.
Kirk PM (1983). New or interesting microfungi IX. Dematiaceous hyphomycetes
from Esher Common. Transactions of the British Mycological Society 80:
449–467.
Kirk PM, Cannon PF, Minter DW, et al. (2008). Ainsworth and Bisby's Dictionary
of the Fungi,10
th
edn. CAB International, Wallingford, UK.
Kirschner R, Pang KL, Jones EBG (2013). Two cheirosporous hyphomycetes
reassessed based on morphological and molecular examination. Mycolog-
ical Progress 12:29–36.
Knapp DG, Kov
acs GM, Zajta E, et al. (2015). Dark septate endophytic pleo-
sporalean genera from semiarid areas. Persoonia 35:87–100.
Kobayasi Y (1971). Mycological reports from New Guinea and the Solomon
Islands (1–11). Bulletin of the National Science Museum (Tokyo) 14:
367–551.
Kodsueb R, Dhanasekaran V, Aptroot A, et al. (2006). The family Pleospor-
aceae: intergeneric relationships and phylogenetic perspectives based on
sequence analyses of partial 28S rDNA. Mycologia 98: 571–583.
Kodsueb R, Lumyong S, Ho WH, et al. (2007). Morphological and molecular
characterization of Aquaticheirospora and phylogenetics of Massarinaceae
(Pleosporales). Botanical Journal of the Linnean Society 155: 283–296.
Kohlmeyer J (1969). Marine fungi of Hawaii including the new genus Helicascus.
Canadian Journal of Botany 47: 1469–1487.
Kohlmeyer J, Volkmann-Kohlmeyer B (1991). Illustrated key to the filamentous
marine fungi. Botanica Marina 34:1–61.
Kohlmeyer J, Volkmann-Kohlmeyer B, Eriksson OE (1995). Fungi on Juncus
roemerianus. 2. New dictyosporous ascomycetes. Botanica Marina 38:
165–174.
Kolomiets T, Pankratova L, Mukhina Z, et al. (2008). First report of leaf spot
caused by Periconia igniaria on yellow starthistle in Russia. Plant Disease
92: 983.
Kruys Å, Wedin M (2009). Phylogenetic relationships and an assessment of
traditionally used taxonomic characters in the Sporormiaceae (Pleosporales,
Dothideomycetes,Ascomycota), utilising multi-gene phylogenies. System-
atics and Biodiversity 7: 465–478.
Lawrey JD, Diederich P, Nelsen MP, et al. (2012). Phylogenetic placement of
lichenicolous Phoma species in the Phaeosphaeriaceae (Pleosporales,
Dothideomycetes). Fungal Diversity 55: 195–213.
Leuchtmann A (1984). Über Phaeosphaeria Miyake und andere bitunicate
Ascomyceten mit mehrfach querseptierten Ascosporen. Sydowia 37:
75–194.
Liew ECY, Aptroot A, Hyde KD (2002). An evaluation of the monophyly of
Massarina based on ribosomal DNA sequences. Mycologia 94: 803–813.
Liu JK, Hyde KD, Jones EBG, et al. (2015). Fungal diversity notes 1–110:
taxonomic and phylogenetic contributions to fungal species. Fungal Diversity
72:1–197.
Liu JK, Phookamsak R, Jones EBG, et al. (2011). Astrosphaeriella is poly-
phyletic with species in Fissuroma gen. nov., and Neoastrosphaeriella gen.
nov. Fungal Diversity 51: 135–154.
Lucas MT (1968). Culture studies on Portuguese species of Leptosphaeria II.
Transactions of the British Mycological Society 51:411–415.
Lumbsch HT, Huhndorf SM (2010). Outline of Ascomycota –2009. Fieldiana
Life and Earth Sciences 1:1–60.
Luttrell ES (1973). Loculoascomycetes. In: (Ainsworth GC, Sparrow FK,
Sussman AS, eds), The fungi, vol IVAAcademic Press, New York: 135 –219.
Manamgoda DS, Cai L, Bahkali AH, et al. (2011). Cochliobolus: an overview and
current status of species. Fungal Diversity 51:3–42.
Manoharachary C, Kunwar IK, Rao NK (2007). Two new species of Dictyo-
sporium from India. Indian Phytopathology 60: 341–344.
Marincowitz S, Crous PW, Groenewald JZ, et al. (2008). Microfungi occurring on
Proteaceae in the fynbos. CBS Biodiversity Series 7:1–166.
Mason EW, Ellis MB (1953). British species of Periconia.Mycological Papers 56:
1–127.
Matsushima T (1971). Microfungi of the Solomon Islands and Papua-New
Guinea. Published by the Author, Kobe, Japan.
McKenzie EHC (2010). Two new dictyosporous hyphomycetes on Rhopalostylis
sapida (Arecaceae) in New Zealand. Mycotaxon 111: 155 –160.
McKenzie EHC, Jones EBG, Hyde KD (eds) (2014). Taxonomy and phylogeny
of Dothideomycetes.Phytotaxa 176:1–323.
Mel'nik VA, Shabunin DA (2010). Rare or rarely revealed fungi Monodictys
capensis and Peyronelina glomerulata.Mikologiya i Fitopatologiya 44:
410–413.
Minnis AM, Kennedy AH, Grenier DB, et al. (2012). Phylogeny and taxonomic
revision of the Planistromellaceae including its coelomycetous anamorphs:
contributions towards a monograph of the genus Kellermania.Persoonia 29:
11–28.
Mouzouras R, Jones EBG (1985). Monodictys pelagica, the anamorph of Ner-
eiospora cristata (Halosphaeriaceae). Canadian Journal of Botany 63:
2444–2447.
Munk A (1953). The system of the Pyrenomycetes. A contribution to a natural
classification of the group Sphaeriales sensu Lindau. Dansk Botanisk Arkiv
15:1–163.
Munk A (1956). On Metasphaeria coccodes (Karst.) Sacc. and other fungi
probably related to Massarina Sacc. Massarinaceae n. fam. Friesia 5:
303–308.
Munk A (1957). Danish Pyrenomycetes. A preliminary flora. Dansk Botanisk
Arkiv 17:1–491.
Nagasawa E, Otani Y (1977). Some species of the bambusicolous ascomycetes
from Japan. I. Reports of the Tottori Mycological Institute 15:38–42.
Nag Raj TR (1989). Genera coelomycetum. XXVI: Amarenographium,Callis-
tospora,Hyalothyridium,Orphanocoela anam.-gen. nov., Scolesporiella, and
Urohendersoniella.Canadian Journal of Botany 67: 3169–3186.
Nannizzi A (1934). Repertorio sistematico dei miceti dell'uomo e degli animali.
Trattato di Micopatologia Umana 4:1–557.
Nelsen MP, Lücking R, Grube M, et al. (2009). Unravelling the phylogenetic
relationships of lichenised fungi in Dothideomyceta.Studies in Mycology 64:
135–144.
Ohm RA, Feau N, Henrissat B, et al. (2012). Diverse lifestyles and strategies of
plant pathogenesis encoded in the genomes of eighteen Dothideomycetes
fungi. PLoS Pathogens 8: e1003037.
Olivier C, Berbee ML, Shoemaker RA, et al. (2000). Molecular phylogenetic
support from ribosomal DNA sequences for origin of Helminthosporium from
Leptosphaeria-like loculoascomycete ancestors. Mycologia 92: 736–746.
O'Neill NR, Farr DF (1996). Miscanthus blight, a new foliar disease of orna-
mental grasses and sugarcane incited by Leptosphaeria sp. and its
anamorphic state Stagonospora sp. Plant Disease 80: 980–987.
Pang KL, Jones EBG, Vrijmoed LLP, et al. (2004). Okeanomyces, a new genus
to accommodate Halosphaeria cucullata (Halosphaeriales,Ascomycota).
Botanical Journal of the Linnean Society 146: 223–229.
TANAKA ET AL.
134
Patil VR, Borse BD (2015). Checklist of freshwater mitosporic fungi of India.
International Journal of Bioassays 4: 4090–4099.
Peever TL (2007). Role of host specificity in the speciation of Ascochyta
pathogens of cool season food legumes. European Journal of Plant Pa-
thology 119:119–126.
Phillips AJL, Alves A, Abdollahzadeh J, et al. (2013). The Botryosphaeriaceae:
genera and species known from culture. Studies in Mycology 76:51–167.
Phookamsak R, Liu JK, Chukeatirote E, et al. (2013). Phylogeny and Morphology
of Leptosphaerulina saccharicola sp. nov. and Pleosphaerulina oryzae and
relationships with Pithomyces.Cryptogamie Mycologie 34: 303–319.
Phookamsak R, Liu JK, McKenzie EHC, et al. (2014). Revision of Phaeos-
phaeriaceae.Fungal Diversity 68: 159–238.
Phookamsak R, Manamgoda DS, Li WJ, et al. (2015). Poaceascoma helicoides
gen et sp. nov., a new genus with scolecospores in Lentitheciaceae.
Cryptogamie Mycologie 36: 225–236.
Photita W, Lumyong P, McKenzie EHC, et al. (2002). A new Dictyosporium
species from Musa acuminata in Thailand. Mycotaxon 82: 415–419.
Pirozynski KA (1972). Microfungi of Tanzania. I. Miscellaneous fungi on oil palm.
II. New hyphomycetes. Mycological Papers 129:1–64.
Prasher IB, Verma RK (2015). Two new species of Dictyosporium from India.
Phytotaxa 204: 193–202.
Pratibha J, Prabhugaonkar A (2015). Multi-gene phylogeny of Pithomyces with
the sexual morph of P. flavus Berk. & Broome. Phytotaxa 218:84–90.
Prihatini R, Boonyuen N, Sivichai S (2008). Phylogenetic evidence that two
submerged-habitat fungal species, Speiropsis pedatospora and Xylomyces
chlamydosporus, belong to the order Jahnulales insertae sedis Dothideo-
mycetes.Microbiology Indonesia 2: 136–140.
Quaedvlieg W, Verkley GJM, Shin HD, et al. (2013). Sizing up Septoria.Studies
in Mycology 75: 307–390.
Raja HA, Stchigel AM, Miller AN, et al. (2007). Hyphomycetes from the Great
Smoky Mountains National Park, including three new species. Fungal Di-
versity 26: 271–286.
Raja HA, Tanaka K, Hirayama K, et al. (2011). Freshwater ascomycetes: two
new species of Lindgomyces (Lindgomycetaceae,Pleosporales,Dothideo-
mycetes) from Japan and USA. Mycologia 103: 1421–1432.
Rayner RW (1970). A mycological colour chart. Commonwealth Mycological
Institute, Kew, Surrey. British Mycological Society.
Rehner SA, Buckley E (2005). A Beauveria phylogeny inferred from nuclear ITS
and EF1-αsequences: evidence for cryptic diversification and links to
Cordyceps teleomorphs. Mycologia 97:84–98.
Rehner SA, Samuels GJ (1994). Taxonomy and phylogeny of Gliocladium
analysed from nuclear large subunit ribosomal DNA sequences. Mycological
Research 98: 625–634.
Ridley GS (1988). New records and species of Loculoascomycetes from New
Zealand. New Zealand Journal of Botany 26: 409–422.
Rouxel T, Balesdent MH (2005). The stem canker (blackleg) fungus, Lep-
tosphaeria maculans, enters the genomic era. Molecular Plant Pathology 6:
225–241.
Saccardo PA (1883). Sylloge Fungorum Omnium hucusque cognitorum, Volume
2. Padova, Italy.
Saccardo PA (1884). Sylloge Fungorum Omnium hucusque cognitorum, Volume
3. Padova, Italy.
Saccardo PA (1886). Sylloge Fungorum Omnium hucusque cognitorum, Volume
4. Padova, Italy.
Samuels GJ (1980). Ascomycetes of New Zealand 1. Ohleria brasiliensis and its
Monodictys anamorph, with notes on taxonomy and systematics of Ohleria
and Monodictys.New Zealand Journal of Botany 18: 515–523.
Scheinpflug H (1958). Untersuchungen über die Gattung Didymosphaeria Fuck.
und einige verwandte Gattungen. Berichte der Schweizerischen Bota-
nischen Gesellschaft 68: 325–385.
Scheuer CH (1991). Massarina tetraploa sp. nov., the teleomorph of Tetraploa
aristata.Mycological Research 95: 126–128.
Schoch CL, Crous PW, Groenewald JZ, et al. (2009). A class-wide phylogenetic
assessment of Dothideomycetes.Studies in Mycology 64:1–15.
Schoch CL, Seifert KA, Huhndorf S, et al. (2012). Nuclear ribosomal internal
transcribed spacer (ITS) region as a universal DNA barcode marker for
Fungi.Proceedings of the National Academy of Sciences of the United
States of America 109: 6241–6246.
Schoch CL, Shoemaker RA, Seifert KA, et al. (2006). A multigene phylogeny of
the Dothideomycetes using four nuclear loci. Mycologia 98: 1041–1052.
Seifert KA, Morgan-Jones G, Gams W, et al. (2011). The genera of Hypho-
mycetes. CBS Biodiversity Series 9:1–997.
Semeniuk G (1983). Association of Trematosphaeria circinans with crown and
root rot of alfalfa in South Dakota. Mycologia 75: 744–747.
Sharma R, Sharma R, Crous PW (2015). Matsushimamyces, a new genus from
soil in Central India. IMA Fungus 6: 337–343.
Shearer CA (1993). A new species of Kirschsteiniothelia (Pleosporales) with an
unusual fissitunicate ascus. Mycologia 85: 963–969.
Shearer CA, Huhndorf SM, Crane JL (1993). Reexamination of eight taxa
originally described in Leptosphaeria on members of Asteraceae.Mycologia
85: 825–834.
Shearer CA, Raja HA (2010). Freshwater ascomycetes database.http://fungi.
life.illinois.edu/ (accessed 11 Feb 2014).
Shearer CA, Raja HA, Miller AN, et al. (2009). The molecular phylogeny of
freshwater Dothideomycetes.Studies in Mycology 64: 145–153.
Shenoy BD, Jeewon R, Wang H, et al. (2010). Sequence data reveals phylo-
genetic affinities of fungal anamorphs Bahusutrabeeja,Diplococcium,
Natarajania,Paliphora,Polyschema,Rattania and Spadicoides.Fungal Di-
versity 44: 161–169.
Shenoy BD, Jeewon R, Wu WP, et al. (2006). Ribosomal and RPB2 DNA
sequence analyses suggest that Sporidesmium and morphologically similar
genera are polyphyletic. Mycological Research 110: 916–928.
Shoemaker RA (1984). Canadian and some extralimital Nodulosphaeria and
Entodesmium species. Canadian Journal of Botany 62: 2730–2753.
Shoemaker RA, Babcock CE (1985). Canadian and some extralimital Para-
phaeosphaeria species. Canadian Journal of Botany 63: 1284–1291.
Shoemaker RA, Babcock CE (1989). Phaeosphaeria. Canadian Journal of
Botany 67: 1500–1599.
Shoemaker RA, Eriksson OE (1967). Paraphaeosphaeria michotii. Canadian
Journal of Botany 45: 1605–1608.
Sivanesan A (1984). The bitunicate ascomycetes and their anamorphs.J.
Cramer, Vaduz.
Sivanesan A (1987). Graminicolous species of Bipolaris,Curvularia,Exser-
ohilum and their teleomorphs. Mycological Papers 158:1–261.
Sivanesan A, Waller JM (1986). Sugarcane diseases. Phytopathological Paper
29:1–88.
Sivichai S, Jones EBG, Hywel-Jones NL (2000). Fungal colonisation of wood in
a freshwater stream at Khao Yai National Park, Thailand. Fungal Diversity 5:
71–88.
Sivichai S, Sri-Indrasutdhi V, Jones EBG (2011). Jahnula aquatica and its
anamorph Xylomyces chlamydosporus on submerged wood in Thailand.
Mycotaxon 116: 137 –142.
Slippers B, Boissin E, Phillips AJL, et al. (2013). Phylogenetic lineages in the
Botryosphaeriales: a systematic and evolutionary framework. Studies in
Mycology 76:31–49.
Subramanian CV (1961). On some species of Cryptostictis.Proceedings of the
National Institute of Science, India Part B Biological Sciences 27: 236–245.
Subramanian CV, Sekar G (1987). Three bitunicate ascomycetes and their tretic
anamorphs. Kavaka 15:87–98.
Suetrong S, Boonyue N, Pang KL, et al. (2011a). A taxonomic revision and
phylogenetic reconstruction of the Jahnulales (Dothideomycetes), and the
new family Manglicolaceae.Fungal Diversity 51: 163–188.
Suetrong S, Hyde KD, Zhang Y, et al. (2011b). Trematosphaeriaceae fam. nov.
(Dothideomycetes,Ascomycota). Cryptogamie Mycologie 32: 343–358.
Suetrong S, Schoch CL, Spatafora JW, et al. (2009). Molecular systematics of
the marine Dothideomycetes.Studies in Mycology 64: 155–173.
Sutton BC (1980). The Coelomycetes. Fungi imperfecti with pycnidia, acervuli
and stromata. Commonwealth Mycological Institute, Kew, Surrey, England.
Sutton BC, Alcorn JL (1974). Neottiosporina. Australian Journal of Botany 22:
517–530.
Tamura K, Stecher G, Peterson D, et al. (2013). MEGA6: molecular evolutionary
genetics analysis version 6.0. Molecular Biology and Evolution 30:
2725–2729.
Tanabe AS (2011). Kakusan4 and Aminosan: two programs for comparing
nonpartitioned, proportional and separate models for combined molecular
phylogenetic analyses of multilocus sequence data. Molecular Ecology
Resources 11: 914–921.
Tanaka K, Harada Y (2003a). Pleosporales in Japan (1): the genus Lophios-
toma.Mycoscience 44:85–96.
Tanaka K, Harada Y (2003b). Pleosporales in Japan (3). The genus Massarina.
Mycoscience 44: 173–185.
Tanaka K, Harada Y (2004). Bambusicolous fungi in Japan (1): four Phaeos-
phaeria species. Mycoscience 45: 377–382.
Tanaka K, Harada Y (2005). Bambusicolous fungi in Japan (6): Katumotoa,a
new genus of phaeosphaeriaceous ascomycetes. Mycoscience 46:
313–318.
Tanaka K, Harada Y, Barr ME (2005a). Bambusicolous fungi in Japan (3): a new
combination, Kalmusia scabrispora.Mycoscience 46:110–113.
REVISION OF THE MASSARINEAE
www.studiesinmycology.org 135
Tanaka K, Hatakeyama S, Harada Y (2004). A new species, Massarina mag-
niarundinacea.Mycotaxon 90: 349–353.
Tanaka K, Hatakeyama S, Harada Y (2005b). Three new freshwater ascomy-
cetes from rivers in Akkeshi, Hokkaido, northern Japan. Mycoscience 46:
287–293.
Tanaka K, Hirayama K, Yonezawa H, et al. (2009). Molecular taxonomy of
bambusicolous fungi: Tetraplosphaeriaceae, a new pleosporalean family with
Tetraploa-like anamorphs. Studies in Mycology 64: 175–209.
Tanaka K, Mel'nik VA, Kamiyama M, et al. (2010). Molecular phylogeny of two
coelomycetous fungal genera with stellate conidia, Prosthemium and
Asterosporium,onFagales trees. Botany 88: 1057–1071.
Taylor JE, Hyde KD (2003). Microfungi of tropical and temperate palms. Fungal
Diversity Press, Hong Kong.
Teng SC (1934). Notes on Sphaeriales from China. Sinensia 4: 359–433.
Teng SC (1936). Additional fungi from China. II. Sinensia 7: 490–527.
Thambugala KM, Ariyawansa HA, Liu ZY, et al. (2014). Towards a natural
classification of Dothideomycetes 6: The genera Dolabra,Placostromella,
Pleosphaerellula,Polysporidiella and Pseudotrichia (Dothideomycetes
incertae sedis). Phytotaxa 176:55–67.
Trakunyingcharoen T, Lombard L, Groenewald JZ, et al. (2014). Mycoparasitic
species of Sphaerellopsis, and allied lichenicolous and other genera. IMA
Fungus 5: 391–414.
Tsui CKM, Berbee ML, Jeewon R, et al. (2006). Molecular phylogeny of Dic-
tyosporium and allied genera inferred from ribosomal DNA. Fungal Diversity
21: 157–166.
Tsui CKM, Hyde KD (2004). Biodiversity of fungi on submerged wood in a
stream and its estuary in the Tai Ho Bay, Hong Kong. Fungal Diversity 15:
205–220.
Tsui CKM, Hyde KD, Fukushima K (2003). Fungi on submerged wood in the
Koito River, Japan. Mycoscience 44:55–59.
Tubaki K (1978). On the Skerman's micromanipulator and microforge (in Jap-
anese). Transactions of the Mycological Society of Japan 19: 237–239.
Tzean SS, Chen JL (1989). Two new species of Dictyosporium from Taiwan.
Mycological Research 92: 497–502.
Van Warmelo KT, Sutton BC (1981). Coelomycetes VII. Stegonsporium.
Mycological Papers 145:1–45.
Vasilyeva LN (1998). Pyrenomycetes and Loculoascomycetes (in Russian). In:
(Azbukina ZM, ed), Lower plants, fungi, and bryophytes of the Russian Far
East, vol. IVNauka, St. Petersburg: 1–419.
Verkley GJM, Dukik K, Renfurm R, et al. (2014). Novel genera and species of
coniothyrium-like fungi in Montagnulaceae (Ascomycota). Persoonia 32:25–51.
Verkley GJM, Mel'nik VA, Shin HD, et al. (2005). Camarographium koreanum
sp. nov., a new coelomycete from Korea. Sydowia 57: 259–266.
Verkley GJM, Quaedvlieg W, Shin HD, et al. (2013). A new approach to species
delimitation in Septoria.Studies in Mycology 75: 213–305.
Voglmayr H, Jaklitsch WM (2011). Molecular data reveal high host specificity in
the phylogenetically isolated genus Massaria (Ascomycota,Massariaceae).
Fungal Diversity 46: 133–170.
Voglmayr H, Jaklitsch WM (2014). Stilbosporaceae resurrected: generic
reclassification and speciation. Persoonia 33:61–82.
Von Arx JA, Müller E (1975). A re-evaluation of the bitunicate ascomycetes with
keys to families and genera. Studies in Mycology 9:1–159.
Wanasinghe DN, Jones EBG, Camporesi E, et al. (2014). An exciting novel
member of Lentitheciaceae in Italy from Clematis vitalba.Cryptogamie
Mycologie 35: 323–337.
Webster J (1955). Graminicolous pyrenomycetes. V. Conidial states of Lep-
tosphaeria michotii,L. microscopica,Pleospora vagans and the perfect state
of Dinemasporium graminum.Transactions of the British Mycological So-
ciety 38: 347–365.
Wehmeyer LE (1961). A world monograph of the genus Pleospora and its
segregates. University of Michigan Press, Michigan, USA.
Wei CT (1950). Notes on Corynespora.Mycological Papers 34:1–10.
White TJ, Bruns T, Lee S, et al. (1990). Amplification and direct sequencing of
fungal ribosomal RNA genes for phylogenetics. In: PCR protocols: a guide to
methods and applications (Innis MA, Gelfand DH, Sninsky JJ, White TJ,
eds). Academic Press, San Diego, California: 315–322.
Whitton RS, McKenzie EHC, Hyde KD (2012). Fungi associated with Pan-
danaceae.Fungal Diversity Research Series 21:1–428.
Wijayawardene NN, Bhat DJ, Hyde KD, et al. (2014a). Camarosporium sensu
stricto in Pleosporinae,Pleosporales with two new species. Phytotaxa 183:
16–26.
Wijayawardene NN, Camporesi E, Bhat DJ, et al. (2014b). Macrodiplodiopsis in
Lophiostomataceae,Pleosporales.Phytotaxa 176: 192–200.
Wijayawardene NN, Crous PW, Kirk PM, et al. (2014c). Naming and outline of
Dothideomycetes –2014 including proposals for the protection or sup-
pression of generic names. Fungal Diversity 69:1–55.
Wijayawardene NN, Hyde KD, Bhat DJ, et al. (2014d). Camarosporium-like
species are polyphyletic in Pleosporales; introducing Paracamarosporium
and Pseudocamarosporium gen. nov. in Montagnulaceae.Cryptogamie
Mycologie 35: 177–198.
Wijayawardene NN, Hyde KD, Bhat DJ, et al. (2015). Additions to brown spored
coelomycetous taxa in Massarinae,Pleosporales: introducing Phragmoca-
marosporium gen. nov. and Suttonomyces gen. nov. Cryptogamie Mycologie
36: 213–224.
Wijayawardene DNN, Mckenzie EHC, Hyde KD (2012). Towards incorporating
anamorphic fungi in a natural classification –checklist and notes for 2011.
Mycosphere 3: 157–228.
Wikee S, Lombard L, Nakashima C, et al. (2013). A phylogenetic re-evaluation
of Phyllosticta (Botryosphaeriales). Studies in Mycology 76:1–29.
Wongsawas M, HongKai W, Hyde KD, et al. (2009). Dictyosporium zhejiangense
sp. nov., a new freshwater anamorphic fungus from China. Cryptogamie
Mycologie 30: 355–362.
Woudenberg JHC, Groenewald JZ, Binder M, et al. (2013). Alternaria redefined.
Studies in Mycology 75: 171–212.
Wu WP, Zhuang WY (2005). Sporidesmium,Endophragmiella and related
genera from China. Fungal Diversity Research Series 15:1–351.
Yuan ZQ, Barr ME (1994). New ascomycetous fungi on bush cinquefoil from
Xinjiang, China. Sydowia 46: 329–337.
Zhang H, Hyde KD, Abdel-Wahab MA, et al. (2013). A modern concept for Heli-
cascus with a Pleurophomopsis-like asexual state. Sydowia 65: 147–166.
Zhang J, Zhou Y, Dou Z, et al. (2015). Helicascus unilocularis sp. nov., a new
freshwater pleosporalean ascomycete from the Caribbean area. Mycological
Progress 14:47.
Zhang TY, Zhao GZ, Zhang XG, et al. (2009a). 26 genera of dematiaceous
dictyosporous hyphomycetes excluding Alternaria (in Chinese). In: Flora
Fungorum Sinicorum 31. Science Press, Beijing.
Zhang Y, Crous PW, Schoch CL, et al. (2012). Pleosporales. Fungal Diversity
53:1–221.
Zhang Y, Fournier J, Bahkali AH, et al. (2011). Inflatispora, a novel lignicolous
genus of Pleosporales from France. Sydowia 63: 287–295.
Zhang Y, Fournier J, Jeewon R, et al. (2008a). Quintaria microsporum sp. nov.,
from a stream in France. Cryptogamie Mycologie 29: 179–182.
Zhang Y, Fournier J, Pointing SB, et al. (2008b). Are Melanomma pulvis-pyrius
and Trematosphaeria pertusa congeneric? Fungal Diversity 33:47–60.
Zhang Y, Liu Y, Zhoh Y, et al. (2014b). Helicascus gallicus sp. nov., a new
freshwater pleosporalean ascomycete from France. Phytotaxa 183:
183–192.
Zhang Y, Schoch CL, Fournier J, et al. (2009b). Multi-locus phylogeny of
Pleosporales: a taxonomic, ecological and evolutionary re-evaluation.
Studies in Mycology 64:85–102.
Zhang Y, Wang HK, Fournier J, et al. (2009c). Towards a phylogenetic clarifi-
cation of Lophiostoma/Massarina and morphologically similar genera in the
Pleosporales.Fungal Diversity 38: 225–251.
Zhang Y, Zhang J, Wang Z, et al. (2014a). Neotypification and phylogeny of
Kalmusia.Phytotaxa 176: 164–173.
Zucconi L, Lunghini D (1997). Studies on Mediterranean hyphomycetes. VI.
Remarks on Bactrodesmium, and B. cubense comb. nov. Mycotaxon 63:
323–327.
TANAKA ET AL.
136
