cryptogamie
mycologie
volume 34 n°4 2013
contents
Bernard DUHEM — Phlebia rhodana sp. nov. and Phlebia jurassica sp.
nov. (Agaricomycotina), two new species from France with a tuberculate
hymenophore. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291-301
Rungiwa PHOOKAMSAK, Jian-Kui LIU, Ekachai CHUKEATIROTE, Eric
H.C. McKENZIE & Kevin D. HYDE — Phylogeny and morphology of Leptosphaerulina saccharicola sp. nov. and Pleosphaerulina oryzae and
relationships with Pithomyces. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303-319
Gothamie WEERAKOON & André APTROOT — Some new lichen species
from Sri Lanka, with a key to the genus Heterodermia in Sri Lanka . . . . . . . 321-328
Sarah EVANS, Ryan W. HANSEN, Heather M. STONE & Mark A.
SCHNEEGURT — Isolation and Characterization of Halotolerant Soil Fungi
from the Great Salt Plains of Oklahoma (USA) . . . . . . . . . . . . . . . . . . . . . 329-341
Li-Song WANG, Xin-Yu WANG & H. Thorsten LUMBSCH — Eight
lecanoroid lichen species new to China . . . . . . . . . . . . . . . . . . . . . . . . . . 343-348
Ke ZHANG, Yan-Ying SU & Lei CAI — An optimized protocol of single
spore isolation for fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349-356
Nalin N. WIJAYAWARDENE, Erio CAMPORESI, Yu SONG, Dong-Qin
DAI, D. Jayarama BHAT, Eric H.C. McKENZIE, Ekachai CHUKEATIROTE,
Vadim A. MEL’NIK, Yong WANG & Kevin D. HYDE — Multi-gene analyses
reveal taxonomic placement of Scolicosporium minkeviciusii in Phaeosphaeriaceae (Pleosporales) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357-366
Slavomir ADAMΩIK, Xavier CARTERET & Bart BUYCK — Type studies on
some Russula species described by C.H. Peck . . . . . . . . . . . . . . . . . . . . . 367-391
Table of volume 34 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 393-394
New taxa proposed in Cryptogamie, Mycologie, 34, 2013. . . . . . . . . . . . 395-396
Cryptogamie, Mycologie, 2013, 34 (4): 357-366
© 2013 Adac. Tous droits réservés
Multi-gene analyses reveal taxonomic placement
of Scolicosporium minkeviciusii in Phaeosphaeriaceae
(Pleosporales)
Nalin N. WIJAYAWARDENE a,b,c, Erio CAMPORESI d, Yu SONG a,
Dong-Qin DAI b,c, D. Jayarama BHAT b,c,e, Eric H.C. McKENZIE f,
Ekachai CHUKEATIROTE b,c, Vadim A. MEL’NIK g,
Yong WANG a* & Kevin D. HYDE b,c
a Department
of Plant Pathology, Agriculture College,
Guizhou University, 550025, P.R. China
c School
b Institute of Excellence in Fungal Research and
of Science, Mae Fah Luang University, Chiang Rai, 57100, Thailand
d A.M.B.
Gruppo Micologico Forlivese “Antonio Cicognani”,
Via Roma 18, Forlì, Italy; A.M.B. Circolo Micologico “Giovanni Carini”,
C.P. 314, Brescia, Italy; Società per gli Studi Naturalistici della Romagna,
C.P. 144, Bagnacavallo (RA), Italy
e 128/1-J,
f Manaaki
Azad Housing Society, Curca, Goa Velha 403108, India
Whenua Landcare Research, Private Bag 92170, Auckland, New Zealand
g Laboratory
of the Systematics and Geography of Fungi,
Komarov Botanical Institute, Russian Academy of Sciences,
Professor Popov Street 2, St. Petersburg, 197376, Russia
Abstract – Scolicosporium minkeviciusii, was newly collected in Italy, and subjected to
morpho-molecular analyses. Morphological characters clearly indicate that this species is a
coelomycete. Combined maximum-likelihood and maximum-parsimony analyses of LSU
and SSU gene sequence data of S. minkeviciusii grouped it in Phaeosphaeriaceae with
Phaeosphaeria nodorum, P. oryzae and Stagonospora foliicola, although the type species of
Scolicosporium, S. macrosporium, which has not been sequenced, is considered to belong in
the family Pleomassariaceae. In this study, we designate an epitype for Scolicosporium
minkeviciusii. The placement of S. macrosporium and Scolicosporium sensu stricto remains
uncertain and further morpho-molecular studies are necessary to confirm the taxonomic
placement of this type species and to delimit this genus.
Asexual states / Classification / Coelomycetous fungi / Phylogeny
* Corresponding author: Yong Wang, E-mail: yongwangbis@yahoo.cn
doi/10.7872/crym.v34.iss2.2013.357
358
N. N. Wijayawardene, E. Camporesi , Y. Song et al.
INTRODUCTION
The term “coelomycete”, introduced by Grove (1919), is used for
asexually reproducing fungi which produce conidia “in a cavity lined by
either fungal tissue, host tissue, or a combination of both” (Kirk et al., 2008).
Approximately 1000 genera of coelomycetes have been described
(Wijayawardene et al., 2012b) but most are not placed in a natural classification
system, i.e. neither being linked to a sexual state nor to have a known familial
affiliation (Wijayawardene et al., 2012a). Most established links of sexual and
asexual ascomycete states are based on co-occurrence of both forms in close
proximity on the host (Spooner & Kirk, 1982), or through culture-based methods
(Hyde et al., 1996; Ramaley & Barr, 1996). Since the introduction of molecular
techniques in fungal studies (White et al., 1990), it has become possible to link
sexual and asexual forms using DNA gene sequence analyses. It has, therefore,
become necessary to re-collect species in order to extract DNA and to carry out
morpho-molecular studies to confirm their natural taxonomic placements
(Wijayawardene et al., 2012c).
The genus Scolicosporium was introduced by Roumeguère (1880) for
S. fagi Lib. ex Roum. Saccardo (1881) synonymized Sporidesmium vermiforme
Riess and Coryneum macrosporium Berk. under Scolicosporium fagi (as
Scolecosporium fagi). Sutton (1977, 1980) designated the older name, Coryneum
macrosporium, as the type of Scolicosporium [as S. macrosporium (Berk.)
B. Sutton] and synonymized Coryneum macrosporium, Scolicosporium fagi and
Sporidesmium vermiforme with this species. In Index Fungorum (2013), there are
13 epithets listed under Scolicosporium, although most of these epithets have
been placed in other genera (Sutton, 1975; Spooner & Kirk, 1982). Only
three species are presently accepted in Scolicosporium, i.e. S. macrosporium
(Sutton, 1977), S. pauciseptatum Constant. (≡ Hendersonia
fusarioides
Sacc.)
.
.
(Constantinescu, 1991) and S. minkeviciusii Treigiene (Treigiene & Mel’nik,
2002).
Scolicosporium has been treated as a coelomycete (Nag Raj & DiCosmo,
1980; Sutton, 1980), but Spooner & Kirk (1982) considered the genus to have
hyphomycetous affinities. Spooner & Kirk (1982) said that Asteromassaria
macrospora (Desm.) Höhn is the sexual state of Scolicosporium macrosporium,
based on association of both species on the same host. This link has not been
proven by molecular analysis and no Scolicosporium species DNA sequences have
so far been deposited in GenBank.
.
We studied a recent specimen of Scolicosporium minkeviciusii (Treigiene
& Mel’nik, 2002) on Quercus pubescens collected in Italy. The aim of this paper
is to epitypify, re-describe and illustrate this fungus. Single conidial culture was
sequenced using LSU and SSU genes to infer the phylogenetic placement of this
species within the Ascomycota.
MATERIALS AND METHODS
Collection and isolation: Conidiomata were observed by hand lens during the
collection of saprobic fungi associated with dead plant material in Buggiana
(Suasia Valley), Italy. These were collected, placed in paper bags and returned to
Taxonomic placement of Scolicosporium minkeviciusii
359
the laboratory. The specimens were observed under a stereoscope. Conidiomata
were removed, placed in a droplet of distilled water on a clean slide, neatly
squashed and examined under a compound microscope to observe the conidial
characters. Single spore isolation was carried out following the method described
in Chomnunti et al. (2011). Germinating conidia were transferred aseptically to
potato dextrose agar (PDA) plates and grown at 18-20°C. Colony colour and
morphological characteristics were assessed after 5 days and 1 week. The
specimens were deposited in the Mae Fah Luang University (MFLU) Herbarium,
Chiang Rai, Thailand. Living cultures were also deposited in Landcare Research,
New Zealand (ICMP), Mae Fah Luang University Culture Collection (MFLUCC)
and Department of Plant Pathology, Agriculture College, Guizhou University,
China (GHUP).
DNA extraction, PCR amplification and sequencing: A Biomiga Inco. kit (GD2416,
San Diego, CA 92121) was used to extract genomic DNA from fresh fungal
mycelia. For the amplification of internal transcribed spacers (ITS), small subunit
nuclear rRNA gene region (SSU) and large subunit nuclear rRNA gene region
(LSU), ITS5 and ITS4, NS1 and NS4 (White et al., 1990) and LROR and LR5
(Vilgalys & Hester, 1990) primers were used respectively. DNA amplification was
carried out as previously described by Liu et al. (2012). The amplified genes were
then sent to SinoGenoMax Co., Beijing, China for DNA sequencing. The
nucleotide sequence data obtained were deposited in GenBank (Table 1).
Phylogenetic analyses: Generated sequences of LSU (KF 366382) and SSU (KF
366383) nuclear rRNA were used for molecular phylogenetic analyses and the
sequence alignments were deposited in TreeBASE (http://www.treebase.org/;
submission ID 14584). A BLAST search in GenBank for LSU and SSU sequences
which followed alignments for Dothideomycetes in Schoch et al. (2006) and Zhang
et al. (2009, 2012) revealed the closest taxa to our strain. Different sequences
from the closest taxa in different families (Coniothyriaceae, Cucurbitariaceae,
Didymellaceae, Leptosphaeriaceae, Phaeosphaeriaceae and Pleosporaceae) were
selected. These sequences were downloaded and aligned separately using Bioedit
(Hall, 2004) and ClustalX (Kohli & Bachhawat, 2003). Alignments were checked
and manual adjustments made where appropriate. Individual datasets were
concatenated into a combined dataset. The ITS gene sequence was excluded in the
phylogenetic analyses, but are deposited in GenBank as it is the preferred locus for
use in fungal barcoding (Schoch et al., 2011).
Maximum likelihood (ML) analysis was performed in RAxML
(Stamatakis, 2006) implemented in raxmlGUI v.0.9b2 (Silvestro & Michalak,
2012). Fifty thorough maximum likelihood (ML) tree searches were done in
RAxML under the general time reversible model (GTR), with each one starting
from a separate randomised tree and the best scoring tree selected. One thousand
non parametric bootstrap iterations were run with the GTR model and a discrete
gamma distribution (Liu et al., 2012).
Maximum-parsimony analysis was carried out using PAUP v. 4.0b10
(Swofford, 2003), and was performed using the heuristic search option with 1000
random taxa additions and tree bisection and reconnection (TBR) as the branch
swapping algorithm. All characters were unordered and of equal weight and gaps
were treated as missing data. Max trees were unlimited, branches of zero length
were collapsed and all multiple, equally parsimonious trees were saved. Clade
stability was assessed using a bootstrap (BT) analysis with 1000 replicates, each
with 10 replicates of random stepwise addition of taxa (Hillis & Bull, 1993). Trees
were visualized with Tree View (Page, 1996).
360
N. N. Wijayawardene, E. Camporesi , Y. Song et al.
Table 1. Sequence data used in this study. Newly produced sequences in bold
GenBank Accession number
Culture collection
number
LSU
SSU
CBS 129.79
EU754128
EU754029
AFTOL-ID 54
AY544645
AY544727
Cochliobolus sativus
AFTOL-ID
DQ678045
DQ677995
Coniothyrium palmarum
CBS 400.71
EU754153
EU754054
Coniothyrium palmarum
CBS 758.73
EU754154
EU754055
Cucurbitaria berberidis
CBS 394.84
GQ387605
GQ387544
Didymella exigua
CBS 183.55
EU754155
EU754056
Didymella pisi
CBS 126.54
GU237968
EU754038
Leptosphaeria doliolum
CBS 541.66
JF740284
Leptosphaeria doliolum
CBS 155.94
JF740282
Leptosphaeria doliolum
CBS 125979
JF740283
Leptosphaeria slovacica
CBS 389.80
JF740315
JF740101
Leptosphaerulina australis
CBS 317.83
GU301830
GU296160
Montagnula anthostomoides
CBS 615.86
GU205223
GU205246
Ophiosphaerella herpotricha
AFTOL-ID 1595
DQ767656
DQ767650
Paraphoma fimeti
CBS 170.70
GQ387584
GQ387523
Peyronellaea zeae-maydis
CBS 588.69
EU754192
EU754093
Phaeosphaeria nodorum
CBS 110109
EU754175
EU754076
Phaeosphaeria oryzae
CBS 110110
GQ387591
GQ387530
Phoma herbarum
CBS 615.75
EU754186
EU754087
Pleospora calvescens
CBS 246.79
EU754131
EU754032
Pleospora herbarum
CBS 191.86
GU238160
GU238232
Pyrenochaeta acicola
CBS 122789
EU754204
EU754204
Pyrenochaeta nobilis
CBS 407.76
EU754206
EU754107
Pyrenochaeta quercina
CBS 115095
GQ387619
GQ387558
Pyrenochaetopsis decipiens
CBS 343.85
GQ387624
GQ387563
Taxon
Ampelomyces quisqualis
Cochliobolus heterostrophus
CBS 101635
GQ387627
GQ387566
AFTOL-ID 283
DQ499596
DQ499595
MFLUCC 12-0089
KF366382
KF366383
Setophoma sacchari
CBS 333.39
GQ387586
GQ387525
Stagonospora foliicola
CBS 110111
EU754217
EU754118
Wojnowicia hirta
CBS 160.73
EU754222
EU754123
MFLUCC 120733
KC594287
KC594288
Pyrenochaetopsis leptospora
Pyrenophora phaeocomes
Scolicosporium minkeviciusii
Wojnowicia viburni
Taxonomic placement of Scolicosporium minkeviciusii
361
RESULTS
Phylogenetic analyses: The combined gene data set of LSU and SSU consists of 32
sequences of 29 taxa with the out group taxon. The dataset consists of
2376 characters including coded alignment gaps; 1660 are constant, while 194 are
variable and 125 are parsimony uninformative in the MP and ML analyses. A best
scoring RAxML tree is shown in Fig. 1. Bootstrap support (BS) values of ML and
MP (equal to or above 50% based on 1000 replicates) are shown above branches.
Our strain of Scolicosporium minkeviciusii (ICMP 19881; MFLUCC 120089) clustered in Phaeosphaeriaeae, however, the phylogenetic relationship of
this species towards the various genera accepted within the family remains
uncertain. The phylogenetic tree also suggested that this strain might be close to
Phaeosphaeria, Setophoma and Stagonospora as they formed a subclade, although
without support.
Fig. 1. RAxML tree generated by the analysis of combined data set of LSU and SSU nrDNA
sequences. Bootstrap support values greater than 50% for maximum likelihood (ML) and
maximum parsimony (MP) analyses are given above the nodes respectively. The epitype strain of
Scolicosporium minkeviciusii (MFLUCC 12-0089) (Phaeosphaeriaceae) is in bold. The tree is
rooted to Montagnula anthostomoides (CBS 615.86) (Montagnulaceae).
362
N. N. Wijayawardene, E. Camporesi , Y. Song et al.
Taxonomy
.
.
Scolicosporium minkeviciusii Treigiene, in Treigiene & Mel’nik, Mikol. Fitopatol.
36(6): 45 (2002)
Figs 2-5
Saprobic on bark of Quercus pubescens Willd. and Q. robur
L. Conidiomata 100–120 µm high × 120–180 µm wide, pycnidial, solitary to
gregarious, elongate-globose, dark brown, uniloculate. Ostiole central, sometimes
towards one side, papillate when young, opening longitudinally at maturity.
Pycnidial wall comprising 3–4 layers, outer layers dark brown, 8–10 µm wide, cells
of textura angularis, inner layers hyaline, 5–40 µm wide. Conidiophores 13–30 ×
2–3 µm, 1–2-septate, branched at the base, hyaline, cylindrical. Conidiogenous
cells annelledic, hyaline, cylindrical. Conidia 60–65 × 7–9 µm (› = 62.2 × 7.7 µm,
n = 20), curved to sigmoid, pale to moderately dark brown, with hyaline end cells,
6–7-transverse eusepta, smooth-walled, tapered to the obtuse apex, base truncate.
Culture characteristics: Conidial germination starts from basal hyaline cell
or first lower median cell. Cells at first swell and produce germ tubes from any
point. The germ tubes may eventually emerge from all cells. On PDA, pale brown
on surface and white at margin, slow growing, attaining a diam. of 1-2 cm after
14 days at 20°C, with thin mycelium, zonate, circular. Reverse of the colony
reddish orange after 2 weeks.
Material examined: Italy, Emilia-Romagna, Forlì-Cesena Province, Santa
Sofia, Buggiana (Suasia Valley), on bark of Quercus pubescens, 19 November
2011, Erio Camporesi, It NNW 40 (MFLU 13–0089, epitype designated here; ex-
Fig. 2. Scolicosporium minkeviciusii on
Quercus pubescens (MFLU 13-0089,
epitype). A. Conidiomata on host. B,
C. Immature conidiomata. D. Mature
sporodochia-like conidiomata. E, F.
Cross sections of immature conidiomata. G. Ostiolar opening. H, I. Wall
of the conidioma. J-M. Different stages
of developing conidia with conidiophores. Scale bars: E 130 µm; F 150 µm;
G 40 µm; H–M 30 µm.
Taxonomic placement of Scolicosporium minkeviciusii
363
epitype culture ICMP 19881 = HGUP N53 = MFLUCC 12–0089); Lithuania,
Vilnius, in silave Tartokiai dicto,
. hab. in ramis emortuis siccis Quercus robur,
15 November 1996, A. Treigiene (LE 212415, holotype).
GenBank accession numbers of ex-epitype: for LSU (KF366382) and
SSU (KF366383).
Fig. 3. Conidia of Scolicosporium minkeviciusii. Scale bars: A-G. 30 µm.
Fig. 4. Scolicosporium minkeviciusii (epitype).
A, B. Germinating conidia. C.
Cultural characters on PDA
from above after
7 days. D. Reverse
culture. Scale bars:
A, B. 30 µm.
364
N. N. Wijayawardene, E. Camporesi , Y. Song et al.
DISCUSSION
There have been only a few studies on Scolicosporium (Saccado, 1881;
.
Sutton, 1975, 1980; Spooner & Kirk, 1982; Constantinescu, 1991; Treigiene &
Mel’nik, 2002) since introduction of the genus by Roumeguère (1880). Several
names have been included in the genus (Index Fungorum, 2013); although Sutton
(1975) concluded that they are not congeneric with S. macrosporium, the type
species of the genus. Sutton (1975) and Spooner & Kirk (1982) excluded several
taxa as they have different morphological characters. These species and the
reasons for their exclusion are listed in Table 2.
Constantinescu (1991) synonymized Hendersonia fusarioides with
S. pauciseptatum and mentioned that three other Scolicosporium species
(S. barringtoniae Viennot-Bourgin, S. gei Chona et al. and S. lactucae Munjal
& Kapoor.) were not discussed by Sutton (1975) and Spooner & Kirk (1982).
Constantinescu (1991) then proceeded to remove these species from
Scolicosporium because, in most cases, conidia
were uniseptate (but
.
S. barringtoniae varies with 1-2-septa). Treigiene & Mel’nik (2002) introduced
S. minkeviciusii, hence there are three accepted species in Scolicosporium i.e.
S. macrosporium, S. barringtoniae and S. minkeviciusii.
Sutton (1975, 1977, 1980) and Nag Raj & DiCosmo (1980) considered
Scolicosporium as coelomycetous, while Spooner & Kirk (1982) discussed
its hyphomycetous affinity. Seifert et al. (2011) listed Scolicosporium as a
hyphomycete. This contrasting understanding is primarily due to the nature of
conidiomata. In S. minkeviciusii, conidiomata are immersed in the substrate when
immature and typically coelomycetous (Fig. 2 B, C). As the conidiomata mature
they become sporodochia-like (Fig. 2 D). The nature of conidioma is very similar
with the definition of acervulus (Kirk et al., 2008). However, the conidioma
of S. minkeviciusii is subglobose and much more typical of a pycnidium than
an acervulus. The longitudinal ostiolar opening found in the conidioma of
Scolicosporium minkeviciusii also agrees with the definition of a pycnidium in
Kirk et al. (2008). Considering these characters, we conclude that S. minkeviciusii
is coelomycetous fungus.
A link between Scolicosporium and its sexual state was reported by
Saccardo (1883) who connected S. macrosporium (as Coryneum macrosporum
Table 2. Species excluded from Scolicosporium
(based on Sutton, 1975 and Spooner & Kirk, 1982)
Species
Sutton (1975)
Spooner & Kirk (1982)
Scolicosporium betulae Rostr.
? Scolicosporium fagi Lib. ex Roum.
S. coryli Dearn. & House
Doubtful
S. fusarioides (Sacc.) B. Sutton
Scolicosporium fusarioides (Sacc.)
B. Sutton
Excipularia fusispora (Berk. &
Broome) Sacc.
Seimatosporium Corda or
Monochaetia (Sacc.) Allesch.
Excipularia fusispora (Berk. &
Broome) Sacc.
S. pedicellatum Dearn. & Overh.
S. phoebes T.S. Ramakr
S. syzygii Ciccar.
S. transversum Fairm.
S. typhae Höhn.
Seiridium Nees
Doubtful
Doubtful
Seiridium Nees
Not mentioned
Seiridium Nees
Doubtful
Doubtful
?Seimatosporium Corda
Scolecosporiella Petr.
Taxonomic placement of Scolicosporium minkeviciusii
365
Berk.) with Asteromassaria macrospora (Desm.) Höhn. (as Massaria macrospora
(Desm.) Sacc.). Höhnel (1917), Spooner & Kirk (1982) and Sivanesan (1984)
accepted this connection as the taxa were observed in close proximity on the same
host. In our analyses of combined LSU and SSU gene regions, Scolicosporium
minkeviciusii groups in Phaeosphaeriaceae (Fig. 1) along with other wellestablished genera, i.e. Ampelomyces, Paraphoma, Phaeosphaeria, Setophoma,
Stagonospora and Wojnowicia (de Gruyter et al., 2009, 2010; Wijayawardene et al.,
2013). We therefore suggest that S. minkeviciusii is placed in Phaeosphaeriaceae,
although better support is needed to resolve relationships with other genera. As
the type species of Scolicosporium, S. macrosporium has not been recollected and
sequenced, its family affinities require further studies.
Acknowledgements. Nalin N. Wijayawardene and D.Q. Dai acknowledge the
Mushroom Research Foundation (MRF), Chiang Rai Province, Thailand for providing
postgraduate scholarship support. Erio Camporesi would like to thank Giancarlo Lombardi
for his invaluable asistance in the collecting programme and identifying host plants.
D. Jayarama Bhat thanks Mae Fah Luang University for a visiting professsorship during
the tenure of which this paper was finalised. Appreciation is extended to the Guizhou
Province Research Fund No. 20113045 for funding all molecular studies and Mae Fah
Luang University grant for studying Dothideomycetes (no. 56101020032).
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