Fungal Diversity (2012) 57:1–44
DOI 10.1007/s13225-012-0175-8
Multigene phylogeny and taxonomy of the genus
Melanconiella (Diaporthales)
Hermann Voglmayr & Amy Y. Rossman &
Lisa A. Castlebury & Walter M. Jaklitsch
Received: 7 March 2012 / Accepted: 15 May 2012 / Published online: 11 July 2012
# Mushroom Research Foundation 2012
Abstract Molecular phylogenetic analyses of LSU rDNA
demonstrate monophyly of the genus Melanconiella, and its
status as a genus distinct from Melanconis is confirmed.
Data of macro- and microscopic morphology, pure cultures
and phylogenetic analyses of partial SSU-ITS-LSU rDNA,
tef1 and rpb2 sequences revealed 13 distinct species of
Melanconiella, six of which are described as new (M. chrysodiscosporina, M. chrysomelanconium, M. chrysorientalis,
M. echinata, M. elegans, M. meridionalis). Melanconiella
hyperopta var. orientalis is described as a new variety.
Diaporthe carpinicola, D. ellisii, D. flavovirens, D. hyperopta and D. ostryae are formally combined into
Melanconiella. The name Melanconiella chrysostroma is
excluded from Melanconiella, as it is an obligate synonym of
Wuestneia xanthostroma. The type of Melanconiella is confirmed as M. spodiaea. Several species are lecto- and/or epitypified. A key to all treated species of Melanconiella is provided,
and the circumscriptions of the genera Melanconis and
Melanconiella are emended. Most Melanconiella species
revealed by molecular phylogenetic analyses can be well characterised by a suite of morphological traits including ascospore
shape, length and width, colour, absence/presence and shape of
appendages and the anamorph. Anamorph-teleomorph connections were confirmed by pure culture and DNA data, revealing
H. Voglmayr (*) : W. M. Jaklitsch
Department of Systematic and Evolutionary Botany,
Faculty Centre of Biodiversity, University of Vienna,
Rennweg 14,
1030, Wien, Austria
e-mail: hermann.voglmayr@univie.ac.at
A. Y. Rossman : L. A. Castlebury
Systematic Mycology & Microbiology Laboratory, USDA,
Agricultural Research Service,
B010A, 10300 Baltimore Ave,
Beltsville, MD 20705, USA
the presence of a single melanconium- or discosporina-like
anamorph for each species. Colony growth was found to be
characteristic of the respective species. Melanconiella is shown
to be confined to the host family Betulaceae, and all species are
found to be highly host-specific, mostly confined to a single host
species. The biodiversity of Melanconiella was determined to be
centred on the genus Carpinus with nine species, five of which
have been confirmed on C. betulus. Europe appears to be the
geographic centre of Melanconiella biodiversity.
Keywords High diversity . Host specificity .
Melanconidaceae . Molecular phylogeny . New species .
Pathogen . Systematics
Introduction
The Diaporthales are an order of ascomycetes within the
Sordariomycetidae that are characterised by forming black
perithecia with or without a well-developed stroma or pseudostroma and a diaporthalean-type centrum development, i.e.
with broad deliquescent paraphyses and asci that float free at
maturity, often having a refractive apical ring (Barr 1978;
Castlebury et al. 2002). Many species occur on hardwood
trees in temperate regions although a few species have been
discovered in subtropical areas. Many species are opportunistic pathogens and endophytes (Sieber 2007), and some of
those are serious threats to forestry and agriculture.
Within the Diaporthales, the family Melanconidaceae
has traditionally been conceived to include many genera
having perithecia immersed in a well-developed stroma with
ostioles (“beaks”) that emerge through an ectostromatic disc
(Barr 1978). Based on phylogenetic data, it was concluded
that the Melanconidaceae should be restricted to a single genus
Melanconis (Castlebury et al. 2002). This narrowly
2
circumscribed family is closely related to the Gnomoniaceae.
Castlebury et al. (2002) determined that species previously
placed in the genus Melanconis are phylogenetically distributed throughout the Diaporthales.
The genus Melanconis, based on the type species M. stilbostoma, includes species with perithecia immersed in the substrate arranged in a circle with oblique or lateral ostioles
convergent and erumpent through an ectostromatic disc and
hyaline ascospores. Melanconis stilbostoma occurs on Betula
and is congeneric with M. alni and M. marginalis, which also
occur on members of the Betulaceae (Castlebury et al. 2002).
The genus Melanconiella Sacc. 1882 was established for
species that are similar to Melanconis but have dark coloured
ascospores, although even Saccardo placed a hyaline-spored
species (M. chrysostroma) in this genus. Subsequently, 21 taxa
were placed in Melanconiella primarily in the late nineteenth
and early twentieth century. Following Wehmeyer (1941), recent authors including Müller and von Arx (1962) and Barr
(1978) considered Melanconiella to be a synonym of
Melanconis, whereas Munk (1957), Petrak (1952a) and
Dennis (1968) preferred to maintain separate genera.
During many years of collecting, the last author noticed a
high variability of ascospore size and morphology, especially
within the hyaline-spored Melanconis species growing on
Carpinus betulus and Ostrya carpinifolia, which all keyed
out as Melanconis (0Melanconiella) chrysostroma based on
the available taxonomic literature (Wehmeyer 1941; Ellis and
Ellis 1997). Results of recent studies on ascomycetes with
similar ecology (e.g. Voglmayr and Jaklitsch 2008, 2011;
Mejía et al. 2011a) revealed a much higher biodiversity than
previously perceived, which cast doubt on the wide species
concept commonly applied to Melanconiella chrysostroma.
The fact that no recent monographic treatment is available for
the genera Melanconis and Melanconiella led us to initiate
thorough morphological and molecular phylogenetic studies
on Melanconiella involving extensive studies of numerous
fresh collections and types from various hosts to evaluate
morphological variability, resolve species boundaries and
clarify host ranges. Cultures were obtained from both ascospores and conidia to elucidate the anamorph-teleomorph
connections by DNA data. In addition, molecular phylogenetic analyses were performed to determine the phylogenetic
relationships within the genus as well as the phylogenetic
position of Melanconiella within the Diaporthales.
Materials and methods
Sample sources
Details of the specimens used for morphological investigations
are listed in the Taxonomy section under the respective descriptions. The herbarium acronyms are given according to Thiers
Fungal Diversity (2012) 57:1–44
(2012): B—Botanischer Garten und Botanisches Museum
Berlin-Dahlem, Germany; BPI—U.S. National Fungus
Collections, Beltsville, USA; DAOM—Agriculture and AgriFood Canada, Ottawa, Canada; G—Conservatoire et Jardin
botaniques de la Ville de Genève, Switzerland; GZU—KarlFranzens-Universität Graz, Austria; K—Royal Botanic Gardens
Kew, UK; M—Botanische Staatssammlung München,
Germany; NY—New York Botanical Garden, USA; UPS—
Uppsala University, Sweden; W—Naturhistorisches Museum
Wien, Austria; WU—Universität Wien, Austria.
Morphology
Morphological observations and measurements were carried
out on a Nikon SMZ 1500 stereo-microscope and after
mounting in tap water or 3 % KOH on a Zeiss
AxioImager A1 compound microscope using Nomarski differential interference contrast (DIC). Images were recorded
with a Nikon DS-Fi1, a Nikon Coolpix 4500 (cultures) or a
Zeiss AxioCam ICc3 digital camera. Measurements are
reported as maxima and minima in parentheses and the
mean plus and minus the standard deviation of a number
of measurements given in parentheses.
Culture preparation, DNA extraction, PCR and sequencing
Single ascospore or conidium isolates were prepared and
grown on 2 % malt extract agar (MEA), or on 2 % corn meal
agar plus 2 %w/v dextrose (CMD) with subsequent transfer
and storage on CM agar slants. To record growth characteristics, they were grown on 2 % MEA at room temperature
under natural daylight.
Mycelium for DNA extraction was grown, freeze-dried
and ground according to Voglmayr and Jaklitsch (2011).
DNA was extracted using the modified CTAB protocol
described in Riethmüller et al. (2002). In collections of
Melanconiella echinata, M. elegans and M. ellisii for which
no cultures could be obtained, DNA was isolated directly
from ascomata or conidiomata according to Voglmayr and
Jaklitsch (2011), but after the chloroform-isoamyl alcohol
step of the CTAB extraction, the supernatant was further
processed with the NucleoSpin Extract II kit (Macherey-Nagel,
Düren, Germany) according to the manufacturer’s instructions
to obtain highly purified DNA.
The partial nuSSU-complete ITS-partial nuLSU rDNA
region was amplified using the primers V9G (de Hoog and
Gerrits van den Ende 1998) and LR5 (Vilgalys and Hester
1990). For amplification of DNA from herbarium specimens,
ITS and LSU were sometimes amplified separately, with the
primers V9G or ITS5 (White et al. 1990) and LR1 for ITS and
with LR0R and LR5 (Vilgalys and Hester 1990) for LSU. For
the type specimen of M. echinata, ITS1 and ITS2 were separately amplified with the primers ITS5/f5.8SR and f5.8SF
Fungal Diversity (2012) 57:1–44
(Jaklitsch and Voglmayr 2011)/LR1, respectively. As the LSU
sequence of M. echinata could not be revealed with the
primers listed above, it was amplified as three separate fragments about 300 bp long, using the primers LR0R/LR2-A (5′
TGCTTTTCATCTTTCGATCAC 3′), LR2R-A (5′
CAGAGACCGATAGCGCAC 3′)/LR3 (Vilgalys and Hester
1990) and LR3R (http://www.biology.duke.edu/fungi/ mycolab/primers.htm)/LR5. A ca. 1.4 kb fragment of the tef1
(translation elongation factor 1 alpha) gene was amplified
with the primers EF1728F (Chaverri and Samuels
2003) and TEF1LLErev (Jaklitsch et al. 2006). The
latter fragment includes the fourth and the fifth intron
and a significant portion of the last large exon. A ca.
1.2 kb fragment of RNA polymerase II subunit B
(rpb2) was amplified using the primer pair fRPB2-5f
and fRPB2-7cr (Liu et al. 1999). PCR products were
purified using an enzymatic PCR cleanup (Werle et al.
1994) as described in Voglmayr and Jaklitsch (2008).
DNA was cycle-sequenced using the ABI PRISM Big
Dye Terminator Cycle Sequencing Ready Reaction Kit
v. 3.1 (Applied Biosystems,Warrington, UK) and the
PCR primers; in addition, primers ITS4 (White et al.
1990) and LR3 were used for the partial nuSSUcomplete ITS-partial nuLSU rDNA region. Sequencing
was performed on an automated DNA sequencer (ABI
3730xl Genetic Analyzer, Applied Biosystems).
Data analysis
To reveal the phylogenetic position of Melanconiella within
the Diaporthales and to investigate whether the genus is
monophyletic, a phylogenetic analysis was performed with
nuLSU rDNA sequences. Sequences of representative species
were selected from Castlebury et al. (2002) and supplemented
with sequences from GenBank; two accessions of
Gaeumannomyces (Magnaporthaceae) were included as outgroup. GenBank accession numbers are given in the phylogenetic tree (Fig. 1). For detailed investigations of species
relationships and delimitation within Melanconiella, a
combined matrix of nuSSU-ITS1-5.8S-ITS2-LSU
rDNA, rpb2 and tef1 sequences was produced for phylogenetic analyses, with Melanconis stilbostoma as outgroup. The GenBank accession numbers of sequences
used in these analyses are given in Table 1.
Sequence alignments for phylogenetic analyses were produced with either MUSCLE version 3.6 (Edgar 2004) or
MAFFT 6.847 (Katoh et al. 2002; Katoh and Toh 2008)
implemented in UGENE 1.10.0 (http://ugene.unipro.ru),
with a maximum of 100 iterative refinements and gap opening
penalties of 1.03 for ITS-LSU and 0.83 for tef1, respectively.
The resulting alignments were checked using BioEdit version
7.0.9.0 (Hall 1999). After the exclusion of excessive leading
and trailing gap regions and of ambiguously aligned positions,
3
the LSU matrix contained 1363 characters. The combined
data matrix contained 4370 characters; viz. 1689 nucleotides
of SSU-ITS-LSU, 1173 nucleotides of rpb2 and 1508 nucleotides of tef1.
In Maximum Likelihood (ML) and Bayesian analyses,
substitution model parameters were calculated separately for
the different gene regions included in the combined analyses. For ML analyses, 500 rounds of random addition of
sequences as well as 500 fast bootstrap replicates were
computed with RAxML (Stamatakis 2006) as implemented
in raxmlGUI 0.95 (Silvestro and Michalak 2011) using the
GTRGAMMA algorithm.
For Bayesian analyses (BA), the GTR + I + G model was
implemented. Bayesian analyses were performed with the
computer program MrBayes (version 3.1.2; Huelsenbeck
and Ronquist 2001). Three parallel runs of four incrementally heated, simultaneous Markov chains were performed
over 5 million generations of which every 500th and 100th
tree was sampled in each run in the LSU and combined
analyses, respectively. The first 1000 and 10000 trees sampled were discarded in the LSU and combined analyses,
respectively, and a 90 % majority rule consensus of the
remaining trees was computed to obtain posterior probabilities. To test convergence of runs, the results were analysed
using AWTY (Nylander et al. 2008); no indication of lack of
convergence was detected.
Maximum parsimony (MP) bootstrap analyses were performed with PAUP* version 4.0 b10 (Swofford 2002), using
1000 replicates of heuristic search with 5 rounds of random
addition of sequences and subsequent TBR branch swapping (MULTREES option in effect, COLLAPSE 0
MAXBRLEN, steepest descent option not in effect); each
bootstrap replicate was limited to 1 million rearrangements.
Results
Molecular phylogenetic analyses
Of the 1363 characters in the LSU alignment, 218 were
parsimony informative. Figure 1 shows the best ML tree
(lnL0−5505.723). Tree topologies of the Bayesian analyses
were fully compatible with the ML analyses. The three
Bayesian runs revealed highly similar posterior probabilities
(PP). ML and MP bootstrap support above 60 % and
Bayesian posterior probabilities above 90 % are given in
Fig. 1 at the first, second and third positions, respectively,
above or below the branches.
Of the 4370 characters in the combined data matrix, 1009
were parsimony informative. Figure 2 shows the best ML
tree (lnL0−16890.279). Tree topologies of the Bayesian
analyses were compatible with the ML tree except for minor
topological differences within some species; in addition, M.
4
Fungal Diversity (2012) 57:1–44
Melanconiella decorahensis MD
Melanconiella flavovirens MFV
-/-/91
Melanconiella carpinicola A.R. 3883
Melanconiella elegans A.R. 3993
Melanconiella meridionalis MOK
Melanconiella ostryae CBS 208.38
95/89/100
Melanconiella chrysomelanconium MCM
74/64/96
98/97/100
98/100/100
Melanconiella chrysodiscosporina MCH
Melanconiella chrysorientalis MGB
Melanconiella echinata DAOM 121196
Melanconiella hyperopta A.R. 3832
92/90/100
Melanconiella spodiaea AF408370
Melanconiella ellisii BPI878343
Phragmoporthe conformis AF408377
-/-/100
Gnomonia gnomon AF408361
77/61/100
87/64/100
Ophiognomonia leptostyla AF408362
Apiognomonia errabunda AF408334
100/99/100
100/100/100
65/-/98
Plagiostoma euphorbiae AF408382
Cryptosporella hypodermia AF408347
75/-/-
Melanconis stilbostoma AF408374
-/-/92
Harknessia eucalypti AF408363
96/70/100
85/75/-
Chromendothia citrina AF408335
Cryphonectria parasitica JN940858
86/92/99
96/97/100
Chrysoporthe cubensis AF408338
Prosthecium pyriforme EU039992
Prosthecium galeatum EU039989
91/82/100
Prosthecium ellipsosporum EU039986
Diaporthe fibrosa AF408351
92/94/100
Diaporthe detrusa AF408349
85/78/100
87/83/100
Diaporthe eres AF408350
Diaporthe pustulata AF408357
Mazzantia napelli AF408368
Valsa ceratosperma AF408386
100/100/100
Leucostoma niveum AF408367
100/100/100
Hercospora tiliae AF408365
99/97/100
Melanconis desmazieri AF408372
97/75/100
81/-/99/100/100
Sillia ferruginea EU683076
Sydowiella fenestrans EU683078
Chapeckia nigrospora EU683068
Hapalocystis berkeleyi AY616230
Pseudovalsa longipes EU683072
Gaeumannomyces medullaris FJ176854
Gaeumannomyces graminis JF414899
0.005 substitutions/site
Fig. 1 Phylogram of the best ML tree (lnL0−5505.723) revealed by
RAxML from an analysis of the LSU matrix of selected Diaporthales,
showing the phylogenetic position of Melanconiella. ML and MP
bootstrap support above 60 % and Bayesian posterior probabilities
above 90 % are given at the first, second and third positions, respectively, above or below the branches. GenBank accession numbers are
given following the taxon names; generic types are formatted in bold
Fungal Diversity (2012) 57:1–44
5
Table 1 Taxa, sources of DNA extracts and GenBank accession numbers of Melanconis and Melanconiella sequences used for the phylogenetic
multi-gene analyses or species identification. Details on the strains are found under specimens examined in the Taxonomy section
Taxon
Strain/culture or specimen
Isolated/extracted from
ITS-LSU
rpb2
tef1
Melanconiella carpinicola
M. carpinicola
M. carpinicola
M. carpinicola
M. carpinicola
M. carpinicola
M. carpinicola
M. chrysodiscosporina
M. chrysodiscosporina
M. chrysodiscosporina
M. chrysodiscosporina
M. chrysodiscosporina
M. chrysodiscosporina
A.R. 3883
MENU
MNM
MNP
MNUK
MSMI
MSR
MCAn
MCH
MCHA
MEE
MEL
MGG
ascospore
conidium
ascospore
ascospore
ascospore
ascospore
ascospore
conidium
ascospore
conidium
conidium
ascospore
ascospore
JQ926230
JQ926231
JQ926232
JQ926233
JQ926234
JQ926235
JQ926236
JQ926237
JQ926238
JQ926239
JQ926240
JQ926241
JQ926242
JQ926303
JQ926304
JQ926305
JQ926306
JQ926307
JQ926308
JQ926309
JQ926310
JQ926311
JQ926312
JQ926313
JQ926314
JQ926369
JQ926370
JQ926371
JQ926372
JQ926373
JQ926374
JQ926375
JQ926376
JQ926377
JQ926378
JQ926379
JQ926380
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
chrysodiscosporina
chrysodiscosporina
chrysodiscosporina
chrysodiscosporina
chrysomelanconium
chrysomelanconium
chrysomelanconium
chrysomelanconium
chrysomelanconium
chrysomelanconium
chrysomelanconium
chrysomelanconium
chrysomelanconium
chrysorientalis
chrysorientalis
chrysorientalis
chrysorientalis
decorahensis
MGJ
MH1
MH2
MRi
MCM
MEH
MEUK
MGH
MGL
MGR
MGS
MGSt
MGUK
MGB
MGP
MGV
MVH
CBS 159.26*
ascospore
ascospore
ascospore
ascospore
ascospore
conidium
conidium
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
conidium
ascospore
JQ926243
JQ926244
JQ926245
JQ926246
JQ926247
JQ926248
JQ926249
JQ926250
JQ926251
JQ926252
JQ926253
JQ926254
JQ926255
JQ926256
JQ926257
JQ926258
JQ926259
JQ926260
JQ926315
JQ926316
JQ926317
JQ926318
JQ926319
JQ926320
JQ926321
JQ926322
JQ926323
JQ926324
JQ926325
JQ926326
JQ926327
JQ926328
JQ926329
JQ926330
JQ926331
JQ926332
JQ926381
JQ926382
JQ926383
JQ926384
JQ926385
JQ926386
JQ926387
JQ926388
JQ926389
JQ926390
JQ926391
JQ926392
JQ926393
JQ926394
JQ926395
JQ926396
JQ926397
JQ926398
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
decorahensis
decorahensis
echinata
elegans
elegans
elegans
elegans
ellisii
ellisii
ellisii
ellisii
flavovirens
flavovirens
flavovirens
MD
MED
DAOM 121196**
CBS 131494*
CBS 131617*
BPI 843574**
BPI 872067**
BPI 843491**
BPI 883227**
BPI 883227**
BPI 878343**
MFA
MFV
MFV1
ascospore
conidium
stroma
ascospore
ascospore
conidioma
conidioma
stroma
conidioma
stroma
stroma
conidium
ascospore
ascospore
JQ926261
JQ926262
JQ926263
JQ926264
JQ926265
JQ926266
JQ926267
JQ926268
JQ926269
JQ926270
JQ926271
JQ926272
JQ926273
JQ926274
JQ926333
JQ926334
JQ926399
JQ926400
JQ926335
JQ926336
JQ926337
JQ926338
JQ926401
JQ926402
JQ926403
JQ926404
JQ926405
JQ926339
JQ926340
JQ926341
JQ926342
JQ926406
JQ926407
JQ926408
JQ926409
MFV2
MFV3
A.R. 3619
ascospore
ascospore
ascospore
JQ926275
JQ926276
JQ926277
JQ926343
JQ926344
JQ926410
JQ926411
M. flavovirens
M. flavovirens
M. hyperopta
6
Fungal Diversity (2012) 57:1–44
Table 1 (continued)
Taxon
Strain/culture or specimen
Isolated/extracted from
ITS-LSU
rpb2
tef1
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
M.
A.R. 3832
CBS 114600*
MCHBV
MCHE
MCL
MCR
MCS
MHG
MHP
MHVA
MSK
MOA
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
conidium
ascospore
conidium
ascospore
conidium
JQ926278
JQ926279
JQ926280
JQ926281
JQ926282
JQ926283
JQ926284
JQ926285
JQ926288
JQ926287
JQ926286
JQ926289
JQ926345
JQ926346
JQ926347
JQ926348
JQ926349
JQ926350
JQ926351
JQ926352
JQ926353
JQ926354
JQ926355
JQ926412
JQ926413
JQ926414
JQ926415
JQ926416
JQ926417
JQ926418
JQ926420
JQ926419
JQ926421
JQ926422
MOK
MOM
MON
MOS
MOSJ
MOSO
MOU
CBS 208.38*
MSH
MVS
SPOD
SPOD1
CBS 121894*
CBS 109778*
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
ascospore
conidium
ascospore
conidium
ascospore
ascospore
JQ926290
JQ926291
JQ926292
JQ926293
JQ926294
JQ926295
JQ926296
JQ926297
JQ926298
JQ926299
JQ926300
JQ926301
JQ926229
AF408374
JQ926356
JQ926357
JQ926358
JQ926359
JQ926360
JQ926361
JQ926362
JQ926363
JQ926364
JQ926365
JQ926366
JQ926367
JQ926302
EU219299
JQ926423
JQ926424
JQ926425
JQ926426
JQ926427
JQ926428
JQ926429
JQ926430
JQ926431
JQ926432
JQ926433
JQ926434
JQ926368
DQ836910
hyperopta
hyperopta
hyperopta
hyperopta
hyperopta
hyperopta
hyperopta
hyperopta
hyperopta var. orientalis
hyperopta var. orientalis
hyperopta var. orientalis
meridionalis
M. meridionalis
M. meridionalis
M. meridionalis
M. meridionalis
M. meridionalis
M. meridionalis
M. meridionalis
M. ostryae
M. spodiaea
M. spodiaea
M. spodiaea
M. spodiaea
Melanconis stilbostoma
M. stilbostoma
* DNA isolated from CBS cultures, ** from herbarium specimens
chrysodiscosporina, M. chrysomelanconium, M. chrysorientalis, M. decorahensis, M. flavovirens, M. meridionalis
and M. ostryae form a monophylum with moderate support
(92–95 % PP) in the Bayesian trees (not shown). The three
Bayesian runs revealed highly similar posterior probabilities. ML bootstrap support above 70 %, MP bootstrap support above 70 % and Bayesian posterior probabilities above
90 % are given in Fig. 2 at the first, second and third
positions, respectively, above or below the branches.
In most cases, ITS sequences were identical or nearly
identical within the species. However, in one accession of
M. chrysodiscosporina (MEL), the ITS showed major differences, and another accession of the same species (MCAn)
had two different ITS copies, but both accessions had rpb2
and tef1 sequences identical to other accessions of M. chrysodiscosporina. Therefore, the ITS sequences of both deviating accessions were removed from the combined matrix
before analysis.
Molecular phylogenetic analyses of the nuLSU matrix
(Fig. 1) confirmed that Melanconiella based on M. spodiaea
is distinct from Melanconis, the latter represented in our
study by its generic type, M. stilbostoma. In addition, monophyly of Melanconiella is highly supported, but its closest
relatives within the Diaporthales remain uncertain due to
partly low backbone support. Internal support is mostly
lacking within Melanconiella in Fig. 1, but the basal position of M. ellisii/M. spodiaea, the position of their neighbour
M. hyperopta and sister group relationship of M. hyperopta
to all other Melanconiella species received high support
(Fig. 1). All 13 Melanconiella species included in the LSU
matrix are revealed as phylogenetically distinct (Fig. 1).
Molecular phylogenetic analyses of the combined matrix
revealed 12 distinct lineages within Melanconiella (Fig. 2),
which are recognised at the species level, and one described
at the varietal level. Of these, Melanconiella spodiaea and
M. decorahensis have already been validly combined; five
lineages represent already described taxa requiring formal
transfer to Melanconiella (Diaporthe carpinicola, D. ellisii,
D. flavovirens, D. hyperopta and D. ostryae); and five
represent yet undescribed species. All species were
Fungal Diversity (2012) 57:1–44
7
As C
G co on Seq
eo sp id u
gr or ium en
.o e
c
Ho rig colocoloe fr
st ins ur ur om
100/75/98
Hosts
Betula pendula
Betula sp.
Carpinus betulus
Carpinus caroliniana
Carpinus orientalis
Corylus avellana
Ostrya carpinifolia
Ostrya virginiana
Geographic origins
A North America
E Europe
Morphology
Ascospores/conidia hyaline
Ascospores/conidia brown
Sequence from
A anamorph
T teleomorph
100/100/100
M. chrysodiscosporina MGJ
M. chrysodiscosporina MH2
M. chrysodiscosporina MCH
M. chrysodiscosporina MGG
M. chrysodiscosporina MEE
M. chrysodiscosporina MRi
100/100/100
M. chrysodiscosporina MH1
M. chrysodiscosporina MCHA
M. chrysodiscosporina MCAn
99/99/100 M. chrysodiscosporina MEL
M. chrysomelanconium MGH
100/99/100
M. chrysomelanconium MGSt
M. chrysomelanconium MGL
M. chrysomelanconium MCM
M. chrysomelanconium MEUK
M. chrysomelanconium MGUK
100/100/100
M. chrysomelanconium MEH
100/100/100 M. chrysomelanconium MGR
M. chrysomelanconium MGS
87/-/- M. chrysorientalis MGP
M. chrysorientalis MGV
M. chrysorientalis MVH
100/100/100 M. chrysorientalis MGB
M. carpinicola MNM
M. carpinicola MSMI
M. carpinicola MNUK
M. carpinicola MNP
100/100/100
M. carpinicola MSR
M. carpinicola MENU
100/-/100
M. elegans CBS 131617
M. elegans BPI 872067
100/100/100
M. elegans CBS 131494
M. elegans BPI 843574
100/100/100 M. decorahensis MD
100/100/100
M. decorahensis MED
M. decorahensis CBS 159.26
M. meridionalis MOA
100/100/100
M. meridionalis MOSO
M. meridionalis MOK
M. meridionalis MOM
72/-/- M. meridionalis MOSJ
100/100/100 M. meridionalis MOU
M. meridionalis MOS
100/100/100
M. meridionalis MON
M. ostryae CBS 208.38
M. flavovirens MFV
93/71/100
M. flavovirens MFV2
100/100/100
M. flavovirens MFV1
M. flavovirens MFA
M. flavovirens MFV3
M. hyperopta MCHBV
M. hyperopta MCR
M. hyperopta MCS
100/100/100 M. hyperopta MCHE
M. hyperopta MHG
M. hyperopta MCL
100/100/100
M. hyperopta var. orientalis MHP
100/100/100
M. hyperopta var. orientalis MHVA
M. hyperopta var. orientalis MSK
M. hyperopta CBS 114600
M. spodiaea MSH
100/100/100
M. spodiaea SPOD
M. spodiaea MVS
99/100/100
98/97/100 M. spodiaea SPOD1
82/72/92 M. ellisii BPI 843491
M. ellisii BPI 878343
100/99/100 M. ellisii BPI 883227
Ms. stilbostoma CBS 121894
Ms. stilbostoma CBS 109778
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
A
A
A
A
E
E
A
E
E
E
E
E
E
E
E
A
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
E
A
A
A
E
E
T
T
T
T
A
T
T
A
A
T
T
T
T
T
A
T
A
T
T
T
T
A
T
T
T
T
T
T
A
T
A
T
A
T
A
T
A
T
T
T
T
T
T
T
T
T
T
T
A
T
T
T
T
T
A
T
T
A
T
T
T
T
A
A
T
T
A
T
T
0.005 substitutions/site
Fig. 2 Phylogram of the best ML tree (lnL0−16890.279) revealed by
RAxML from an analysis of the combined ITS-LSU-rpb2-tef1 matrix
of Melanconiella, with Melanconis stilbostoma selected as outgroup.
ML and MP bootstrap support above 70 % and Bayesian posterior
probabilities above 90 % are given at the first, second and third
positions, respectively, above or below the branches. Strain/culture or
herbarium specimen are given following the taxon names; holo- or
epitype strains/specimens are formatted in bold
8
molecularly highly distinct and received high internal support. The clade containing M. ellisii and M. spodiaea was
placed as sister to the other species with medium (MP) to
high (ML, BA) support (Fig. 2), and within the latter, M.
hyperopta was sister clade to the residual lineages with high
support. Also the two species of Melanconiella on Ostrya
(M. meridionalis, M. ostryae) form a highly supported
clade. Melanconiella chrysodiscosporina, M. chrysomelanconium and M. chrysorientalis, which are highly similar in
their stroma and ascospore features, consistently formed a
highly supported monophylum in nuLSU and combined
analyses, and relationships within that clade also received
high support.
Fungal Diversity (2012) 57:1–44
Chrysostromae of his subgenus Eumelanconis, except for
M. carthusiana, M. desmazierii, M. juglandis, M. pallida
and M. platystroma, which do not belong here. Wehmeyer’s
(1941) sect. Stilbostomae corresponds to Melanconis sensu
stricto, although M. apocrypta, M. antarctica and M. fennica
may not belong there. Melanconis everhartii has been recognized in Macrodiaporthe. Other subgenera that
Wehmeyer (1941) recognized in Melanconis have been
transferred or relocated to other genera, such as Chapeckia
(Barr 1978), Hapalocystis (Jaklitsch and Voglmayr 2004),
Macrodiaporthe, Massariovalsa (Barr 1978), Prosthecium
(Voglmayr and Jaklitsch 2008), Pseudovalsa (M. modonia,
De Silva et al. 2009) and Pseudovalsella (von Höhnel
1918).
Anamorph-teleomorph relationships and cultures
Molecular phylogeny, species delimitation and barcoding
For each Melanconiella species, a single characteristic anamorph could be identified by DNA data obtained from
ascospore and conidial cultures or directly from ascomata
and conidiomata, apart from M. echinata for which no
anamorph is known. Both dark brown melanconium-like
conidia as well as hyaline discosporina-like conidia were
present within the genus Melanconiella, but both were never
found within the same species; conidial morphology was
constant within a species. The production of dark brown and
hyaline conidia did not correlate with phylogenetic relationships, but in several cases, phylogenetic species pairs with
one having hyaline, the other dark brown conidia were
revealed (M. chrysodiscosporina/M. chrysomelanconium;
M. meridionalis/M. ostryae).
Culture characteristics like growth rates, morphological
patterns of colonies on MEA and pigment formation were
observed to be highly distinct for each species (Fig. 3).
Conidiation was not observed on MEA cultures, but one
PDA culture of M. elegans produced aberrant conidia of
variable shapes, the size of which was within the range
observed on the natural substrate.
Discussion
Circumscription of the genus Melanconiella
Morphologically Melanconiella is similar to Melanconis.
Species of the latter genus cause more conspicuous bumps
in the bark of the hosts than those of Melanconiella and
form light-coloured, white or yellowish ectostromatic discs.
Wehmeyer (1941) used this trait to distinguish his section
Stilbostomae from his Chrysostromae, which are characterized by dark coloured discs. Light coloured discs are not
uncommon in Melanconiella, as exemplified by e.g. M.
hyperopta or M. meridionalis. Melanconiella as circumscribed here basically matches Wehmeyer’s (1941) section
Extensive morphological and molecular phylogenetic data for
Melanconiella reveal high species diversity within this genus.
Most of the phylogenetically distinct species of Melanconiella
show a suite of teleomorph characteristics enabling species
identification based on morphology. In some cases, teleomorph features do not allow distinction of closely related
species, but host association as well as anamorphteleomorph relationships provide important information for
reliable identification. As in many host-specific ascomycetes
inhabiting recently dead corticated branches, knowledge of
the host is crucial for correct species identification.
At present the genus Melanconiella does not appear to
belong to any established family within the Diaporthales.
Phylogenetic analyses remain inconclusive about its relationships within the order (Fig. 1). Although previously placed as
a synonym of Melanconis in the Melanconidaceae, this genus
and thus the family have been shown to be closely related to
the Gnomoniaceae (Fig. 1).
Within the genus Melanconiella, all terminal branches
are highly supported, however, the infrageneric relationships remain often unresolved due to lack of backbone
support (Fig. 2). The short branch lengths in the phylograms
indicate that speciation occurred within a short time frame,
obscuring the evolutionary relationships.
The ITS region has been proposed as the primary barcoding locus for fungi (Schoch et al. 2012). In most accessions
of Melanconiella sequenced in our study, the ITS enabled
unequivocal identification. However, in M. chrysodiscosporina we found a deviating sequence in one accession
and two different ITS copies in another accession, whereas
both accessions had rpb2 and tef1 sequences identical to
other accessions of M. chrysodiscosporina. This variation
indicates that the ITS region is not always suitable for
reliable species identification, and results of ITS barcoding
have to be treated with caution. In addition, it also demonstrates that a polyphasic approach is necessary in species-
Fungal Diversity (2012) 57:1–44
9
a
b
c
d
e
f
g
h
i
j
k
l
Fig. 3 Melanconiella cultures on MEA at room temperature, all after
20 d except where noted. a. M. carpinicola (MSMI). b. M. chrysodiscosporina (MGJ). c. M. chrysomelanconium (MEH, 28 d). d. M.
chrysorientalis (MGB). e. M. decorahensis (MED). f. M. elegans
(AR3830, 25 d). g. M. flavovirens (MFV3). h. M. flavovirens (MFA,
63 d). i. M. hyperopta (MCL). j. M. hyperopta var. orientalis (MHP, 31
d). k. M. meridionalis (MOM, 65 d). l. M. spodiaea (MSH, 28 d)
10
Table 2 List of Melanconiella
species, their hosts and distribution. Taxa in bold represent new
species, varieties or
combinations
Fungal Diversity (2012) 57:1–44
Host species
Melanconiella species
Distribution
Betula spp.
C. betulus
C. betulus
C. betulus
C. betulus
C. betulus, C. orientalis
C. caroliniana
C. caroliniana
C. caroliniana
C. orientalis
C. orientalis
Corylus avellana
Ostrya carpinifolia
O. virginiana
M. decorahensis
M. carpinicola comb. nov.
M. chrysodiscosporina sp. nov.
M. chrysomelanconium sp. nov.
M. hyperopta comb. nov.
M. spodiaea
M. echinata sp. nov.
M. elegans sp. nov.
M. ellisii comb. nov.
M. chrysorientalis sp. nov.
M. hyperopta var. orientalis var. nov.
M. flavovirens comb. nov.
M. meridionalis sp. nov.
M. ostryae comb. nov.
Europe, North America
Europe
Europe
Europe
Europe
Europe
North America
North America
North America
Europe
Europe
Europe, ?North America
Europe
Europe
rich groups for robust and reliable identification, and it is
important to sequence also other genes (e.g. Gazis et al.
2011). ITS alone is definitely not sufficient to provide
reliable characterisation of closely related species in many
ascomycete lineages (e.g. Voglmayr & Jaklitsch 2008;
Jaklitsch 2009, 2011; Pavlic et al. 2009; Druzhinina et al.
2010; Jaklitsch et al. 2012).
Biodiversity and nomenclature of Melanconiella
on Carpinus
Until now, only two species of Melanconiella (usually as
Melanconis) have been commonly recognised on Carpinus:
the brown-spored M. spodiaea and the hyaline-spored M.
chrysostroma. This was mainly influenced by Wehmeyer
(1941), who was subsequently widely followed, e.g. by Ellis
and Ellis (1997). Wehmeyer (1941) listed numerous synonyms for M. chrysostroma, however, he may not have examined all of the numerous authentic or type specimens he cited,
as re-examination of several type specimens revealed that they
are not even diaporthalean (see below). According to the spore
size (14–23×3.5–7 μm) listed in Wehmeyer (1941), his concept of M. chrysostroma was apparently based on a mixture of
M. carpinicola and M. hyperopta. Wehmeyer (1941) also
established Melanconis chrysostroma var. ellisii for North
American collections from Carpinus, which were considered
to differ mainly by smaller ascospores.
Present investigations show that M. chrysostroma cannot
be retained, because it is an obligate synonym of Wuestneia
xanthostroma (see below). The molecular phylogenetic investigations show that specimens referred to as M. chrysostroma
are highly heterogeneous, consisting of at least eight species.
Three of these taxa (M. carpinicola, M. ellisii, M. hyperopta)
have been described in the 19th century with accurate descriptions, but had been synonymised under M. chrysostroma.
Anamorphs
Our pure culture and DNA sequence data revealed the presence of a single, characteristic anamorph for each species of
Melanconiella, which is either melanconium- or discosporinalike. This contrasts with Wehmeyer (1937, 1941), who
recorded at least two different types of conidia (alpha and beta)
for several species now placed in Melanconiella. In his pure
culture studies of M. decorahensis, M. ellisii and M. ostryae, he
reported the production of only a single anamorph, and his
conclusions on alpha- and beta-conidia were mainly based on
associations found in herbarium specimens. Knowing that
several species of Melanconiella can co-occur on the same
branch, it is not surprising that this observational evidence
resulted in recording different conidial forms for one species.
Several of the anamorphs of Melanconiella have been
described as species of Melanconium. According to Sutton
(1980), more than 200 binomials have been described in
Melanconium, and no generic revision is available. As many
of the Melanconium binomials date back to the early 19th
century, they cannot be interpreted based on the scanty original descriptions, which often indicate a mixture of discordant
elements based on the hosts listed. Original material is often
not extant, in bad condition, or unavailable for examination. In
addition, the few morphological characters and the similarity
of many Melanconium species are serious obstacles for appropriate interpretation. Most Melanconium names have not
been used in the last decades. Therefore, clear preference
should be given to the teleomorph epithets, which are mostly
well interpretable, as well as to the respective teleomorph
genera that were historically associated with Melanconium.
In addition to Melanconium, species of the anamorph genera
Discosporium and Discosporina have been linked to teleomorphs now placed in Melanconiella. According to Sutton
(1980), the teleomorph of the type species of Discosporium,
Fungal Diversity (2012) 57:1–44
D. populeum (Sacc.) Sutton, is Cryptodiaporthe populea
(Sacc.) Butin (now Plagiostoma populinum (Fuckel) L.C.
Mejía; Mejía et al. 2011b). All Discosporium species attributable to Melanconiella have been transferred to Discosporina
(Petrak 1962); of the four species placed in Discosporina, three
(D. carpinicola and D. deplanata from Carpinus; D. sulphurea
from Corylus) are anamorphs of Melanconiella, but only D.
sulphurea can be unequivocally attributed to a species, viz. M.
flavovirens, the latter having priority.
Ecology and host range
Like many Diaporthales inhabiting woody plants, all species
of Melanconiella are restricted to recently dead corticated
branches of their hosts, often colonising extensive parts. This
is indicative of weak pathogenicity or opportunistic growth,
which appears to be common within the Diaporthales. As the
Diaporthales have been shown to be common endophytes in
broadleaf trees, often remaining dormant until host tissue
senescence (Sieber 2007), species of Melanconiella may be
widely present in healthy branches. This is supported by the
rapid and extensive development of Melanconiella anamorphs
commonly observed on recently cut branches.
All species of Melanconiella were observed to be highly
host-specific as they were found to be confined mostly to a
single, sometimes a few closely related hosts (Table 2). It is not
surprising that we found a new species and variety on
Carpinus orientalis (M. chrysorientalis, M. hyperopta var.
orientalis) and a new species on Ostrya carpinifolia (M. meridionalis), hosts that are largely understudied. Whereas most
host genera of Melanconiella harbour only a single or two
species that do not co-occur in the same area, the situation is
markedly different with Carpinus, which harbours the vast
majority of Melanconiella biodiversity. On Carpinus betulus,
several species of Melanconiella can co-occur in close vicinity,
sometimes even on the same branches, which can pose problems for identification. However, there appears to be some
ecological differentiation. Some species were only found on
11
branches still attached to the tree in open habitats (e.g. M.
carpinicola, M. chrysomelanconium), whereas others mostly
occurred on recently fallen or cut branches within forests (M.
hyperopta, M. chrysodiscosporina), or were concentrated on
branches still attached to the trees within forests (M. spodiaea).
Some of the species also show different seasonality. While M.
hyperopta has mainly been found in the winter season from
late autumn to spring, M. carpinicola, M. chrysodiscosporina
and M. chrysomelanconium were mostly found in the summer
season from late spring to autumn. The reasons for this evolutionary radiation on a single host genus are unknown, but
similar cases are known for other corticolous ascomycetes,
e.g. Prosthecium (Voglmayr and Jaklitsch 2008) or Massaria
(Voglmayr and Jaklitsch 2011), where biodiversity is concentrated on the host genus Acer. In those cases, co-occurrence of
several related species on the same host has been observed,
indicating that this may be a widespread phenomenon in
weakly pathogenic corticolous ascomycetes.
Geographic distribution
Inhabiting hosts belonging to the temperate family Betulaceae,
the genus Melanconiella is confined to the north temperate
zone. Remarkably, the diversity of Melanconiella is highest in
Europe with currently nine species and one variety, followed
by North America with five confirmed species. The report of
M. spodiaea on Castanea crenata from Japan by Kobayashi
(1970) is likely based on a misidentification. The only additional Asian record referable to Melanconiella has been recently published by Vasilyeva and Stephenson (2010), who report
M. ellisii (as M. carpinigera) from the Russian Far East.
Considering that several taxa were included in M. ellisii until
the current study, detailed morphological and DNA studies are
necessary to reveal its true identity. There is little species
overlap between Europe and North America; only M. decorahensis has been confirmed for both continents in the present
study, whereas conspecificity of North American and European
accessions of M. flavovirens remains to be verified.
Taxonomy
Key to the genera Melanconis and Melanconiella
Ectostroma conspicuous, typically projecting up to 1 mm above the substrate, light coloured, white, cream or yellowish; conidia
brown .............................................................................................................................................................. Melanconis
Ectostroma inconspicuous, typically projecting up to 0.5 mm above the substrate, light or dark coloured; conidia hyaline or
brown ............................................................................................................................................ Melanconiella
Melanconis Tul. & C. Tul., Selecta fung carpol (Paris) 2:115
(1863), emend.
Type: Melanconis stilbostoma (Fr.) Tul., Ann Sci nat, ser
4, 5:109 (1856).
Pseudostromata consisting of a well-developed, conspicuous, distinctly erumpent, light coloured, white or yellow
ectostromatic disc causing a coarsely pustulate bark surface.
Ectostromatic disc surrounded by bark or not. Central
12
column beneath the disc more or less conical, comprising
hyaline hyphae mixed with a hyaline or light-coloured powdery amorphous substance; residual entostroma between perithecia of scant hyaline hyphae. Stromatic zones lacking.
Perithecia often appearing as rounded bumps beneath the bark
surface surrounding the ectostromatic disc, oblique or horizontal, often arranged in a circle around the central column,
with long lateral ostioles that converge at the margin of the
column and emerge at the margin or within the ectostromatic
disc, flat discoid or slightly projecting and convex, typically
not obscuring the disc. Paraphyses broad bands, deliquescent
at maturity. Asci oblong or fusoid, octosporous, with an apical
ring typically distinct when fresh; asci becoming detached
from their base. Ascospores hyaline, ellipsoid, bicellular, with
a central or slightly eccentric, not or slightly constricted septum, smooth, with or without blunt or pointed appendages.
Conidial state: melanconium-like. Conidiomata acervular; typically possessing the same type of ectostromatic disc
and central column as the teleomorph, usually preceding it.
Conidiophores branched only at the base, septate, fewcelled, smooth, hyaline. Conidiogenous cells annellidic.
Conidia brown, ellipsoid or subglobose.
Melanconiella Sacc., Syll Fung 1:741 (1882), emend.
Type: Melanconiella spodiaea (Tul.) Sacc., Syll Fung
1:741 (1882); lectotype designated by Müller and von
Arx (1962).
Pseudostromata consisting of an inconspicuous, erumpent, light or more frequently dark coloured ectostromatic
disc or pulvillus (referred to as ectostromatic disc below)
causing a more or less pustulate bark surface. Ectostromatic
disc convex, flat or concave, variously coloured, surrounded
by bark or not. Central column beneath the disc more or less
conical, comprising hyaline or pigmented hyphae mixed
with a pigmented, most commonly cream, yellow, olive,
brownish or gray, powdery amorphous substance; residual
entostroma between perithecia mostly of scant hyphae.
Stromatic zones lacking. Perithecia inconspicuous at the
bark level or appearing as rounded bumps beneath the bark
surface surrounding the ectostromatic disc, oblique or horizontal, scattered or often arranged in a circle around the
central column, with long lateral ostioles that converge at
the margin of the central column or within. Ostioles emerging in various positions in the ectostromatic disc, flat in the
disc or slightly projecting, rarely distinctly projecting and
cylindrical, often obscuring the disc. Paraphyses broad
bands, deliquescent at maturity. Asci oblong or fusoid,
octosporous, with a distinct apical ring, becoming detached
from their base. Ascospores hyaline, yellowish or brown,
fusoid or ellipsoid, bicellular, with a central or slightly
eccentric, not or slightly constricted septum, smooth; wall
Fungal Diversity (2012) 57:1–44
swelling in water mounts or not, with or without short, blunt
appendages and sometimes with a narrow gelatinous sheath.
Conidial state: melanconium- or discosporina-like.
Conidiomata acervular, peridermal or subperidermal; possessing the same type of ectostromatic disc and central
column or lighter than that of the teleomorph, usually preceding it. Conidiophores branched and septate only at the base,
few-celled, smooth, hyaline to light brown. Conidiogenous cells
annellidic (in melanconium-like conidiation) or phialidic (in
discosporina-like conidiation). Conidia brown or hyaline, ellipsoid, subglobose, ovoid or oblong, with or without distinct
hyaline sheath.
Notes: In the original description of Melanconiella,
Saccardo (1882) mentioned uniseptate ascospores becoming
brown at maturity and he listed two species, M. spodiaea
and M. chrysostroma, without selecting a generic type.
Although Clements and Shear (1931) lectotypified the genus with M. chrysostroma, this species is in serious conflict
with the generic protologue, as it is an obligate synonym of
Wuestneia xanthostroma (see below), a taxon of the
Diaporthales with one-celled, hyaline ascospores. The lectotypification of Clements and Shear (1931) is therefore
superseded according to the ICBN Art. 9.17(b). We agree
with Müller and von Arx (1962) who considered M. spodiaea to be the type species of Melanconiella and thus the
lectotypification is attributed to them.
Anamorphs of Melanconis and Melanconiella are usually
referred to as Melanconium. Although Melanconium Link,
Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 9 (1809) is older
than the teleomorph genera, it is not used here. Its type species
M. atrum is said to occur on Fagus sylvatica (Sutton 1980),
but intense searches for many years has not yielded any
Melanconium on Fagus. Re-examination of the lectotype of
M. atrum (K(M) 171588) revealed conidia with a lighter
brown equatorial zone and a size of 10–13×6–8 μm, which
do not fit known anamorphs of Melanconiella. Conidial size
and morphology of M. atrum is similar to the anamorph of
Melanconis alni, which, however, does not occur on Fagus.
Identity, systematic affiliation and phylogenetic position of
Melanconium atrum are therefore obscure.
Melanconiella carpinicola (Fuckel) Voglmayr & Jaklitsch,
comb. nov. (Fig. 4)
MycoBank MB 800113
Basionym: Diaporthe carpinicola Fuckel, Jb Nassau Ver
Naturk 27–28:37 (1874) [1873–74].
Synonym: Diaporthe kunzeana Sacc., Nuovo Giorn Bot
Ital 8:180 (1876) (type not seen).
Pseudostromata 0.8–1.7 mm diam, inconspicuous,
slightly projecting or not, perithecia only rarely projecting;
centre bluntly conical. Ectostromatic disc circular or oblong,
Fungal Diversity (2012) 57:1–44
13
a
b
d
f
e
j
i
c
l
k
r
s
t
g
h
m
n
o
p
q
u
v
w
x
y
c1
z
d1
a1
e1
f1
g1
b1
h1
i1
j1
Fig. 4 Melanconiella carpinicola. a. Panoramic view of pseudostromata. b. Pseudostroma in transverse section showing central column
and perithecia immersed in the entostroma. c. Pseudostroma showing
perithecial bumps, ectostromatic disc and ostioles in surface view. d–f.
Ectostromatic discs and ostioles in surface view. g, h. Transverse
section below ectostromatic disc. i. Ascus. j. Ascus tip with apical
ring. k–y. Ascospores. z. Conidioma in surface view. a1. Transverse
k1
l1
m1
section of conidioma, showing central column. b1, c1. Conidiophores.
d1–m1. Conidia. k, l in 3 % KOH, all other mounts in water. a02 mm,
b, c, z, a10500 μm, d–h0300 μm, i–y, b1–m1010 μm. Sources: a, m.
WU 31806; b, c, f, h. WU 31802; d, g, k, l. G 00111395 (lectotype); i,
j, n–s. WU 31801; t–y, c1, f1, g1, h1. WU 31805; z–b1, i1–m1. WU
31807; d1, e1. M-0177882 (isotype)
14
cream, pale yellowish to dark brown or black, 0.15–0.6 mm
long, sometimes concealed by densely arranged ostioles,
often pulvinate; bark often darkened in a ring around the
disc. Central column grey. Entostroma indistinct. Ostioles
1–10 per disc, 40–125 μm diam, plane or slightly papillate,
black. Perithecia 0.3–0.5 mm diam, arranged in various
configurations. Asci fusoid, (60–)67–82(–92)×8–11(–13)
μm (n029), containing 8 irregularly biseriate ascospores,
with apical ring distinct when fresh, 2.5–2.8 μm diam, 1.3–
1.8 μm high, only faintly visible in older herbarium specimens. Ascospores hyaline, fusoid, distinctly constricted at
the septum, (12–)15–18(–22)×(3.0–)3.8–5.3(–6.5), l/w0
(2.5–)3.1–4.3(–6.2) (n0388) μm; without appendages; cells
dimorphic, upper cell mostly larger, with subacute end,
lower cell narrowly rounded, multiguttulate containing
mostly one large and numerous small guttules per cell; wall
ca. 0.2–0.4 μm thick, not swelling.
Anamorph: discosporina-like. Conidiomata 1–2.5 mm
diam, visible as darker spots, mostly with central or eccentric stromatic column; at maturity covered by whitish discharged conidial masses. Conidiogenous cells phialidic, 10–
18×1.5–2.5 μm, hyaline. Conidia hyaline, elongate to suballantoid, (10–)12–14(–15.5)×(2.7–)3.2–3.7(–4.0) μm, l/
w0(2.8–)3.3–4.2(–5.1) (n0115), multiguttulate; wall ca.
0.2–0.4 μm, without gelatinous sheath.
Culture: Colony on MEA after 17 d at room temperature
ca. 55 mm diam, without a distinct radial arrangement of
surface hyphae, first white, turning dull yellow, eventually
dark brown; aerial hyphae frequent only in the centre; diffusing pigment lacking.
Habitat and host range: dead corticated twigs and
branches of Carpinus betulus attached to the tree; it has
only been found in open sites like forest edges and hedges.
Distribution: Europe; uncommon but widely distributed.
Typification: GERMANY, Hessen, near Oestrich, on dry
branches of Carpinus betulus, autumn, L. Fuckel, Fungi
Rhenani 2660 (G 00111395, lectotype of Diaporthe carpinicola here designated; M-0177882, M-0177883 and WU:
isotypes). AUSTRIA, Niederösterreich, Michelbach Markt,
Mayerhöfen, map grid 7860/4, on corticated branches of
Carpinus betulus, 18 June 2011, H. Voglmayr & W.
Jaklitsch, strain MSMI (from teleomorph) (WU 31801, epitype of Diaporthe carpinicola here designated; ex-epitype
culture CBS 131706), ex-epitype sequences JQ926235
(ITS-LSU), JQ926307 (rpb2), JQ926373 (tef1).
Additional specimens examined: AUSTRIA, Kärnten, St.
Margareten im Rosental, Kramerstrauch, map grid 9452/4, 12
May 2010, H. Voglmayr, strain MNM (from teleomorph) (WU
31802). ibid., 6 July 2002, W. Jaklitsch, strain A.R. 3883 (from
teleomorph) (BPI 841338). ibid., Gupf, near Brici (Writze), 6
July 2006, W. Jaklitsch, W.J. 2924 (WU 31803). ibid.,
Trieblach, near Ledra, soc. M. chrysomelanconium, 7 June
1996, W. Jaklitsch, W.J. 891 (WU 31804). Oberösterreich,
Fungal Diversity (2012) 57:1–44
Raab, map grid 7647/2, soc. M. chrysomelanconium, 1 Aug.
2010, H. Voglmayr, strain MSR (from teleomorph) (WU
31805). GERMANY, Sachsen-Anhalt, Eisleben, Oct. 1875,
G. Winter, Thümen, Mycoth. Univ. 469 (WU). SLOVENIA,
Gorenjska, Povodje, 16 May 2010, H. Voglmayr & W.
Jaklitsch, strain MNP (from teleomorph) (WU 31806).
SWEDEN, Skåne, Äsphult par., Forshult, 8 July 1978, K. &
L. Holm (UPS 412042a, as Melanconis chrysostroma).
UNITED KINGDOM, West Yorkshire, Wakefield,
Newmillerdam N Barnsley, Newmillerdam Country Park,
soc. M. chrysomelanconium, 17 May 2011, H. Voglmayr &
W. Jaklitsch, strains MNUK (from teleomorph), MENU (from
anamorph) (WU 31807, culture CBS 131700).
Notes: Melanconiella carpinicola is well characterised by
protruding and narrowly aggregated ostioles mostly at the
margin of a small inconspicuous ectostromatic disc, which,
at low magnification, gives them a typical appearance of
distinct, small, elevated, black bumps on the twigs. In addition, its small ascospores without swelling walls in combination with its growth on Carpinus betulus are distinctive.
Ecologically, this species is apparently confined to open
habitats, where it is commonly associated with M. chrysomelanconium even on the same twigs. When co-occurring,
the areas of both species can be easily distinguished by their
abundant conidiomata which produce hyaline conidia in M.
carpinicola and dark brown conidia in M. chrysomelanconium. The description of conidia by Fuckel (1874) is in
agreement with our data; the anamorph-teleomorph connection was proven in the present study by DNA data from
conidial and ascospore cultures.
Considering the preservation and abundance of the collection, the part of Fuckel’s original collection of Herbier
Barbey-Boissier housed in G is designated as lectotype here.
To ensure nomenclatural stability, a recent collection, for
which a culture and sequences are available, is designated
here as epitype. Although no type specimen could be located and investigated, Diaporthe kunzeana is clearly a synonym of M. carpinicola according to the general description,
particularly the ascospore size (16–19×3.5–4 μm) and host
given in Saccardo (1876).
Melanconiella chrysodiscosporina Voglmayr & Jaklitsch,
sp. nov. (Fig. 5)
MycoBank MB 800114
Etymology: referring to the yellow stroma and the
discosporina-like anamorph.
Pseudostromata typically indistinct, ill-defined, confluent, irregular, sometimes with circular outline, 1.5–3.4 mm
diam, typically projecting by less than 0.5 mm; perithecial
bumps sometimes distinct. Ectostromatic disc minute, 0.2–
0.6(–0.8) mm long, circular when young, then usually
Fungal Diversity (2012) 57:1–44
15
a
b
d
i
f
e
j
k
q
c
g
m
l
r
s
h
n
t
u
o
p
v
w
a1
x
b1
y
c1
d1
e1
z
f1
g1
h1
Fig. 5 Melanconiella chrysodiscosporina. a. Panoramic view of pseudostromata. b. Pseudostroma in transverse section. c. Three pseudostromata in surface view. d–g. Ectostromatic discs and ostioles in
surface view. h. Transverse section below ectostromatic disc. i. Ascus.
j. Ascus tip with apical ring. k–w. Ascospores (k, w showing
appendages). x. Conidioma in surface view. y. Transverse section of
conidioma, showing central column. z, a1. Conidiophores. b1–h1.
Conidia. All mounts in water. a02 mm, b, c, x, y0500 μm, d–h0
300 μm, i–w, z–h1010 μm. Sources: a, e. WU 31861; b–d, x, y, b1–f1.
WU 31867; f, g. WU 31864; h, z, a1. WU 31862; g1, h1. WU 31824
narrowly fusoid-oblong-linear, often filled by ostioles, light
or bright yellow or greyish to shades of brown. Central
column yellow, greyish yellow, yellow-olive or (light)
olive. Entostroma crumbly, of subhyaline to yellowish
hyphae. Ostioles 1–8(–10) per disc, 50–160 μm diam,
variously arranged in the disc, black, short papillate,
16
apically rounded or flat, sometimes acute with tips 15–
30 μm wide, only slightly projecting above the disc
level. Perithecia 0.3–0.8 mm. Asci cylindrical to fusoid,
(90–)100–119(–127)×(10–)11–15(–18) μm (n039), containing 8 uni- or irregularly biseriate ascospores, with
distinct apical ring, 3.0–3.8 μm diam, 2.0–3.3 μm high.
Ascospores hyaline, broadly ellipsoid, not constricted at
the septum, (15.5–)17–20(–22) × (6.2–)7.2–10.0(–14.5)
μm, l/w 0(1.4–)1.9–2.4(–2.9) (n 0276); ends broadly
rounded to truncate, with hyaline cap-like appendages
1–2.5 μm long, 2.2–5.2 μm wide, fading and eventually
absent in microscope mounts; cells monomorphic, multiguttulate with one large and numerous small guttules per
cell; wall distinctly swelling and sometimes distinctly
bulging particularly at the septum, 0.6–1.4 μm thick,
becoming ca. 1.2–4.7 μm thick at the septum.
Anamorph: discosporina-like. Conidiomata 0.7–4.7 mm
diam, visible as often lobed, darker spots lined by a distinct
dark brown to blackish marginal zone, with a central or
eccentric stromatic column; at maturity covered by whitish
discharged conidial masses. Conidiogenous cells phialidic,
18–32×2.5–4.5 μm, hyaline. Conidia hyaline, ellipsoid,
oblong or cylindrical, (12–)13–15.5(–19)×(4.3–)4.8–5.5(–
6.5) μm, l/w0(2.1–)2.4–3.2(–4) (n0227), with two large
and numerous small guttules; wall ca. 0.4 μm thick, with
distinct gelatinous sheath ca. 0.6–0.8 μm thick.
Culture: Colony on MEA after 17 d at room temperature
ca. 56 mm diam, not zonate, surface sometimes appearing
mottled due to varying mycelial density, aerial hyphae irregularly arranged, abundant only in the centre or causing a
finely floccose surface; first whitish or pale yellowish, turning brown from the centre, margin turning dull yellowbrown or black; diffusing pigment lacking; odour indistinct.
Habitat and host range: dead corticated twigs and
branches of Carpinus betulus mostly lying on the ground.
Distribution: confined to Europe where it is common.
Type: AUSTRIA, Niederösterreich, Michelbach Markt,
Mayerhöfen, map grid 7860/4, on corticated branches of
Carpinus betulus, 20 June 2009, W. Jaklitsch & H.
Voglmayr, strain MCH (from teleomorph) (WU 31859, holotype; ex-holotype culture CBS 125597), ex-holotype sequences JQ926238 (ITS-LSU), JQ926310 (rpb2), JQ926376 (tef1).
Additional specimens examined (all on Carpinus betulus): AUSTRIA, Kärnten, St. Margareten im Rosental, map
grid 9452/4, 23 Sep. 2010, H. Voglmayr & W. Jaklitsch
(WU 31860). Niederösterreich, Gießhübl, Seewiese, map
grid 7863/3, 6 June 2010, H. Voglmayr, strain MGG (from
teleomorph) (WU 31861). Gießhübl, Wassergspreng, map
grid 7963/1, 15 Aug. 2010, H. Voglmayr (WU 31862).
Hardegg, Merkersdorf, Semmelfeld, map grid 7161/3,
8 Aug. 2010, W. Jaklitsch & H. Voglmayr (WU 31863).
Mannersdorf am Leithagebirge, Schweingraben, map grid
8065/2, soc. M. hyperopta (teleomorph), 17 April 2010, H.
Fungal Diversity (2012) 57:1–44
Voglmayr & I. Greilhuber, strain MCAn (from anamorph)
(WU 31824). Michelbach Markt, Mayerhöfen, map grid
7860/4, soc. M. chrysomelanconium, 20 June 2009, W.
Jaklitsch & H. Voglmayr, strain MCH (WU 31864). ibid.,
3 June 2000, W. Jaklitsch, W.J. 1464 (BPI 747940, WU
31872). Mödling, Hinterbrühl, map grid 7963/2, 20 June
2010, H. Voglmayr (WU 31865). Oberösterreich, Raab,
Riedlhof, map grid 7647/2, 4 April 2010, H. Voglmayr,
strain MRi (from teleomorph) (WU 31866). Steegen,
Rittberg, map grid 7648/3, 28 June 2010, H. Voglmayr,
strain MEE (from anamorph) (WU 31867). Wien, Döbling,
Hermannskogel, map grid 7763/2, 18 April 2010, H.
Voglmayr, strains MH1, MH2 (from teleomorph), MCHA
(from anamorph) (WU 31868). Favoriten, Laaerbergstraße,
map grid 7864/3, 8 April 2010, W. Jaklitsch & H. Voglmayr,
strain MEL (from teleomorph) (WU 31869). Floridsdorf,
Jedlersdorferstraße/Marchfeldkanalweg, map grid 7764/1,
soc. M. chrysomelanconium, 9 May 2010, W. Jaklitsch,
strain MGJ (from teleomorph) (WU 31870, culture CBS
131703). Liesing, Maurerwald, map grid 7863/1, soc. M.
chrysomelanconium, 10 July 1993, W. Jaklitsch (WU
15980). CZECH REPUBLIC, Moravia, Lednice, palace
grounds, 1 Oct. 2010, W. Jaklitsch & H. Voglmayr (WU
31871). SWEDEN, Skåne, Äsphult par., Forshult, 8 July
1978, K. & L. Holm (UPS 412042b, as Melanconis
chrysostroma).
Notes: Melanconiella chrysodiscosporina is similar to the
two closely related species M. chrysomelanconium and M.
chrysorientalis, differing mainly in ITS, rpb2 and tef1
sequences; in addition M. chrysomelanconium has dark
brown conidia and slightly larger ascospores than those of
M. chrysodiscosporina and M. chrysorientalis occurs on a
different host, Carpinus orientalis. Melanconiella chrysodiscosporina is most commonly found on recently cut
branches lying on the ground, where it is often associated
with M. hyperopta. As M. chrysodiscosporina has sometimes been found to co-occur on the same branches with the
highly similar M. chrysomelanconium, reliable species identification is only possible on anamorph traits. The
anamorph-teleomorph connection was repeatedly proven
by DNA data from conidial and ascospore cultures.
Melanconiella chrysomelanconium Voglmayr & Jaklitsch,
sp. nov. (Fig. 6)
MycoBank MB 800115
Etymology: referring to the yellow stroma and the
melanconium-like anamorph.
Pseudostromata 1.5–2.6 mm diam, more or less circular,
typically prominent, usually causing distinct bumps in the
bark, but often densely aggregated or confluent, projecting
to ca. 0.5 mm; individual perithecial contours causing small
bumps in the bark. Ectostromatic disc small, 0.2–0.7(–0.9)
Fungal Diversity (2012) 57:1–44
17
a
b
d
i
c
f
e
g
h
j
k
l
m
n
o
p
q
r
s
t
u
x
v
y
w
z
a1
b1
Fig. 6 Melanconiella chrysomelanconium. a. Panoramic view of
pseudostromata. b. Pseudostroma in transverse section. c. Pseudostroma showing ectostromatic disc and ostioles in surface view. d–g.
Ectostromatic discs and ostioles in surface view. h. Transverse section
below ectostromatic disc. i. Ascus. j. Ascus tip with apical ring. k–u.
Ascospores (k–n, s, u showing appendages). v. Conidioma in surface
c1
view. w. Transverse section of conidioma (left), showing central column, and of young stroma and perithecium (right). x. Conidiophores.
y–c1. Conidia. All mounts in water. a02 mm, b, c, v, w0500 μm, d–
h0300 μm, i–u, x–c1010 μm. Sources: a, d, e, j. WU 31878; b, i, p–s.
WU 31874; c, x–c1. WU 31879; f–h, o, w. WU 31875; k, l. WU
31873; m, n. WU 31876; t, u. WU 15180; v. WU 31877
18
mm, usually fusoid or oblong, sometimes circular, yellow,
yellow-brown, greyish yellow or light to dark brown. Central
column often crumbly, bright yellow, grey, brown, olive.
Entostroma inconspicuous or crumbly and olive. Ostioles 1–
10 (commonly up to 6) per disc, 50–160 μm diam at their base,
ca. 30–65 μm at their tips, variably located in the disc, papillate
or with flat apex, brown or shiny black. Perithecia 0.4–0.8 mm
diam; connection to the central column often difficult to ascertain. Asci cylindrical, (93–)109–128(–133)×(12–)14–19(–22)
μm (n022), containing 8 uni- or irregularly biseriate ascospores, with distinct apical ring, 3.5–4.5 μm diam, 2.4–3 μm
high. Ascospores hyaline, broadly ellipsoid, not constricted at
the septum, (16–)19–23(–27.5)×(7.5–)9.0–12.3(–15.5) μm,
l/w0(1.1–)1.8–2.2(–2.6) (n0307); ends broadly rounded to
truncate, with hyaline cap-like appendages 1.7–3 μm long,
3–5.4 μm wide, fading and disappearing in microscope
mounts; cells monomorphic, multiguttulate with one large
and numerous small guttules per cell; wall 0.6–2.6 μm thick,
distinctly swelling particularly at the septum, sometimes distinctly bulging out becoming 1.5–4.8 μm thick at the septum.
Anamorph: melanconium-like. Conidiomata 0.6–4 mm
diam, visible as dark brown to blackish spots, with central or
eccentric stromatic column; at maturity covered by black discharged conidial masses. Conidiogenous cells annellidic, 18–
40×3–5 μm, hyaline. Conidia dark brown, broadly ellipsoid to
broadly pip-shaped, (12.5–)13–15(–16)×(8.5–)8.8–9.5(–10.0)
μm, l/w0(1.3–)1.4–1.6(–1.8) (n098), multiguttulate with 1–2
large and numerous small guttules; wall ca. 0.6–0.8 μm thick,
with distinct gelatinous sheath 0.8–1.1(–1.4) μm thick.
Culture: Colony on MEA after 17 d at room temperature
ca. 56 mm diam, after 4 wk plate nearly entirely covered by
mycelium. Colony circular, with diffuse concentric zones;
aerial hyphae short, only frequent or abundant in the centre,
forming diffuse, transparent whitish zones, below these
zones colony pale yellowish-greenish, turning dark brown,
grey to nearly black from the centre; margin dull golden
yellow to olive; odour slightly sour; diffusing pigment lacking, agar faintly yellowish.
Habitat and host range: dead corticated twigs and
branches of Carpinus betulus attached to the tree; it has
only been found in open sites like forest edges, parks and
hedges.
Distribution: Europe; uncommon but widely distributed.
Type: AUSTRIA, Wien, Floridsdorf, Marchfeldkanalweg,
map grid 7764/1, on corticated branches of Carpinus betulus,
9 Nov. 2008, W. Jaklitsch, strain MCM (from teleomorph)
(WU 31873, holotype; ex-holotype culture CBS 124271), exholotype sequences JQ926247 (ITS-LSU), JQ926319 (rpb2),
JQ926385 (tef1).
Additional specimens examined (all on Carpinus betulus):
AUSTRIA, Kärnten, St. Margareten im Rosental, Trieblach,
near Ledra, map grid 9452/4, soc. M. carpinicola, 7 June 1996,
W. Jaklitsch, W.J. 891 (WU 31804). Niederösterreich,
Fungal Diversity (2012) 57:1–44
Michelbach Markt, Mayerhöfen, map grid 7860/4, soc. M.
chrysodiscosporina, 20 June 2009, W. Jaklitsch & H.
Voglmayr (WU 31864). Oberösterreich, Raab, map grid
7647/2, soc. M. carpinicola, 1 Aug. 2010, H. Voglmayr, strain
MGR (from teleomorph) (WU 31877). Wien, Favoriten,
Laaerbergstraße/Fontanastraße, map grid 7864/3, 30 May
1993, W. Jaklitsch (WU 15180). Floridsdorf, Lazarsfeldgasse,
map grid 7764/2, 23 May 2010, W. Jaklitsch, strain MGL
(from teleomorph) (WU 31874). Floridsdorf,
Jedlersdorferstraße, map grid 7764/1, soc. M. chrysodiscosporina, 9 May 2010, W. Jaklitsch (WU 31870). Landstraße,
Botanical Garden, map grid 7864/1, 23 May 2010, H.
Voglmayr, strains MGH (from teleomorph), MEH (from anamorph) (WU 31875, culture CBS 131699). Landstraße,
Stadtpark, map grid 7764/3, 3 Sep. 2010, H. Voglmayr & W.
Jaklitsch, strain MGSt (from teleomorph) (WU 31876).
Liesing, Maurerwald, map grid 7863/1, soc. M. chrysodiscosporina, 10 July 1993, W. Jaklitsch (WU 15980). SPAIN,
Asturias, Luarca, finca ‘Fonte Baixa’, 8 May 2010, E. Rubio
Domínguez, E.R.D. 5108, strain MGS (from teleomorph) (WU
31878). UNITED KINGDOM, West Yorkshire, Wakefield,
Newmillerdam N Barnsley, Newmillerdam Country Park,
soc. M. carpinicola, 17 May 2011, H. Voglmayr & W.
Jaklitsch, strains MGUK (from teleomorph), MEUK (from
anamorph) (WU 31879, culture CBS 131701).
Notes: Melanconiella chrysomelanconium is similar to
the closely related M. chrysodiscosporina and M. chrysorientalis, from which it differs mainly by dark brown conidia,
slightly larger ascospores and ITS, rpb2 and tef1 sequences;
M. chrysorientalis differs also by the host (Carpinus orientalis). Although M. chrysodiscosporina and M. chrysomelanconium share the same host, they appear to be
ecologically separated: Melanconiella chrysodiscosporina
has been mainly collected on fallen or cut branches within
forests, whereas M. chrysomelanconium is apparently confined to open habitats, where it is commonly associated with
M. carpinicola on twigs attached to the tree. However, M.
chrysomelanconium has sometimes also been found to cooccur on the same branches with M. chrysodiscosporina at the
edges of larger forests. Therefore, reliable identification is
only possible with the anamorph. The anamorph-teleomorph
connection was repeatedly proven by DNA data from conidial
and ascospore cultures. The teleomorph of the Spanish collection is beautifully illustrated with macro- and microphotographs at http://www.ascofrance.com/search_recolte/2696 or
http://www.asturnatura.com/fotografia/setas-hongos/melan
conis-chrysostroma-fr-tul-c-tul-3/9039.html.
Melanconiella chrysorientalis Voglmayr & Jaklitsch, sp.
nov. (Fig. 7)
MycoBank MB 800116
Etymology: referring to the yellow stroma and its host,
Carpinus orientalis.
Fungal Diversity (2012) 57:1–44
19
a
b
d
e
j
i
k
r
s
z
d1
f
l
m
t
u
a1
e1
f1
c
h
g
n
o
v
p
w
x
h1
Fig. 7 Melanconiella chrysorientalis. a. Panoramic view of pseudostromata. b. Pseudostroma in transverse section. c. Three pseudostromata in surface view. d–g. Ectostromatic discs and ostioles in surface
view. h. Transverse section below ectostromatic disc. i. Ascus. j. Ascus
tip with apical ring. k–y. Ascospores (k–m showing appendages). z.
i1
y
c1
b1
g1
q
j1
k1
l1
m1
Conidioma in surface view. a1. Transverse section of conidioma,
showing central column. b1, c1. Conidiophores. d1–m1. Conidia. All
mounts in water. a02 mm, b, c, z, a10500 μm, d–h0300 μm, i–y, b1–
m1010 μm. Sources: a, b, e–h, k–m, z–m1. WU 31858; c, d. WU
31880; i, j, n–y. WU 31881
20
Pseudostromata inconspicuous, 0.9–2.4 mm diam,
slightly (0.1–0.3 mm) projecting, circular or contours illdefined, merging; perithecial contours often evident.
Ectostromatic discs 0.1–0.6 mm, often densely disposed,
with oblong, elliptic or circular outline, yellowish, light
yellow, dull yellow, dark yellow-brown or grey. Central
column crumbly, dull yellow, greenish yellow or olive;
entostroma of light or olive-green hyphae. Ostioles 1–10
per disc, 50–140 μm diam, shiny black or brown with black
centre, convex-papillate, scarcely projecting. Perithecia
0.3–0.6 mm diam; connection to central column often unclear. Asci cylindrical, (104–)114–130(–137)×(11–)13–15
(–17) μm (n033), containing 8 uni-, rarely biseriate ascospores, with distinct apical ring, 3.8–4.3 μm diam, 2.5–
3.2 μm high. Ascospores broadly ellipsoid, not constricted
at the septum, (15.7–)17–19(–22)× (6.0–)7.5–9.3(–10.2)
μm, l/w0(1.7–)1.9–2.4(–3) (n0111); ends broadly rounded
to sometimes truncate, with hyaline cap-like appendages 1.3–
2 μm long, 2.4–4.2 μm wide, fading and eventually absent in
microscope mounts; cells monomorphic, multiguttulate, each
with one large and numerous small guttules; wall distinctly
swelling and sometimes distinctly bulging out particularly at
the septum, 0.6–1.1 μm thick, 1.3–3 μm at the septum.
Anamorph: discosporina-like. Conidiomata 1–2 mm
diam, only scarcely visible in face view, with diffuse margins, with central or eccentric stromatic column; at maturity
covered by discharged whitish to greyish conidial masses.
Conidiogenous cells phialidic, 9–18×2.5–4 μm, hyaline.
Conidia hyaline, ellipsoid, oblong or cylindrical, (11.7–)
12.3–14.0(–15.3)×(5.5–)6.0–6.8(–7.3) μm, l/w0(1.7–)1.9–
2.2(–2.7) (n079), with 1–2 large and numerous small guttules; wall ca. 0.5 μm thick, with distinct gelatinous sheath
ca. 0.6–0.9 μm thick.
Culture: Colony on MEA after 17 d at room temperature
ca. 61 mm diam, after 4 wk mycelium (nearly) covering the
entire plate. Colony forming several concentric zones, each
covered by a conspicuous, whitish, stellate mat of aerial
hyphae; brown zones progressing from the centre, eventually dark brown below the whitish structure; agar pale
yellow to brownish; odour sour.
Habitat and host range: dead corticated twigs and
branches of Carpinus orientalis; only found on branches
attached to the trees.
Distribution: South-eastern Europe, uncommon; collected in Croatia and Greece (Corfu) but likely to co-occur with
its host in other countries.
Type: CROATIA, Istria, Barbariga, on corticated branches
of Carpinus orientalis, 15 May 2010, H. Voglmayr & W.
Jaklitsch, strain MGB (from teleomorph) (WU 31880, holotype; ex-holotype culture CBS 131702), ex-holotype sequences JQ926256 (ITS-LSU), JQ926328 (rpb2), JQ926394 (tef1).
Additional specimens examined (all on Carpinus orientalis): CROATIA, Istria, Poreč, Gedici, 1 Nov. 2010, H.
Fungal Diversity (2012) 57:1–44
Voglmayr & I. Greilhuber, strain MGP (from teleomorph)
(WU 31881). Vrsar, soc. M. spodiaea (anamorph), 14 May
2010, H. Voglmayr & W. Jaklitsch, strains MGV (from
teleomorph), MVH (from anamorph) (WU 31858).
GREECE, Kerkyra (Corfu), W Analipsis, 24 April 2012,
H. Voglmayr & W. Jaklitsch (WU 32000). SE Ano
Korakiana, 24 April 2012, H. Voglmayr & W. Jaklitsch
(WU 32001). S Kouramaditika, 22 April 2012, H.
Voglmayr & W. Jaklitsch (WU 32002).
Notes: Melanconiella chrysorientalis is similar to the
closely related M. chrysodiscosporina and M. chrysomelanconium, from which it differs mainly by ITS, rpb2 and tef1
sequences and a different host (C. orientalis) and from M.
chrysomelanconium by hyaline conidia. The anamorphteleomorph connection was proven by DNA data from conidial and ascospore cultures.
Melanconiella decorahensis (Ellis) Sacc., Syll Fung 2:
Addenda Vol I:LIV (1883). (Fig. 8)
MycoBank MB 210729
Basionym: Melanconis decorahensis Ellis, [as
‘decoraensis’], Am Nat 17:195 (1883).
Synonyms: Melanconis decorahensis var. subviridis Ellis
& Everh., North American Pyrenomycetes 528 (1892) (type
not seen).
Melanconiella subviridis (Ellis) Dearn. & House, Bull
NY State Mus 266:81 (1925).
Pseudostromata 1.5–3 mm diam, circular, often inconspicuous, with their centre typically projecting to ca. 0.6,
rarely 0.8 mm; perithecial contours only rarely evident.
Ectostromatic discs variable, mostly narrow, fusoid or linear, 0.2–1.6 mm long, pale yellow, bright yellow, greenish
yellow, greenish grey, olive or dull greyish brown. Central
column grey, greenish grey or olive-brown. Entostroma
inconspicuous. Ostioles variously arranged in the disc, 1–
20 per disc, 40–140(–160) μm diam, shiny black, flat or
papillate, tips typically flattened. Perithecia 0.3–0.5 mm
diam. Asci narrowly cylindrical, (114–)125–145(–150)×
(11–)12–14(–16) μm (n 035), containing 8 uniseriate
ascospores, with an apical ring distinct when fresh,
3.7–4.2 μm diam, 3.7–4 μm high, often indistinct in
older herbarium specimens. Ascospores dark brown, ellipsoid, distinctly constricted at the septum, (13.5–)16–
20(–22)×(7.5–)8.5–10(–11) μm, l/w0(1.6–)1.8–2.2(–2.4)
(n 057); ends broadly rounded, without appendages;
cells more or less monomorphic, multiguttulate; wall
not swelling, ca. 0.7–0.8 μm.
Anamorph: melanconium-like. Conidiomata 1.5–2.5 mm
diam, blackish brown, at maturity covered by black discharged
conidial masses, forming conspicuous thick black pustules or
spots 1–3 mm diam, sometimes confluent to 1–2 cm.
Conidiogenous cells distinctly annellidic, 20–35×3.5–4 μm,
Fungal Diversity (2012) 57:1–44
21
a
b
d
e
f
k
l
m
r
s
t
c
g
n
u
i
h
o
v
p
q
w
x
a1
j
z
y
Fig. 8 Melanconiella decorahensis. a. Panoramic view of pseudostromata. b. Pseudostroma in transverse section. c. Pseudostroma in surface view. d–f. Ectostromatic discs and ostioles in surface view. g–i.
Transverse sections below ectostromatic disc. j. Ascus. k. Ascus tip
with apical ring. l–x. Ascospores. y. Conidioma in surface view. z, a1.
b1
c1
d1
e1
f1
g1
h1
i1
Conidiophores. b1–i1. Conidia. v–x in 3 % KOH, all other mounts in
water. a02 mm, b–g, y0500 μm, h, i0200 μm, j–x, z–i1010 μm.
Sources: a–c, f–h, j–u, y–e1. WU 31808; i. Wehmeyer 3644 (NY); v–x,
f1–i1. Ellis & Everhart, N. Amer. Fungi. Ser. II: 1562 (NY)
22
at first hyaline, becoming brown with age. Conidia dark
brown, pip-shaped, (13–)15–17(–19)×(8–)9–10(–11) μm, l/
w0(1.2–)1.5–2(–2.3) (n0121), multiguttulate when fresh,
with a large guttule when dead; wall ca. 0.6–0.7 μm thick,
with hyaline gelatinous sheath ca. 0.5 μm thick.
Culture: Colony on MEA after 17 d at room temperature
ca. 67 mm diam, whitish, outline irregular, covering the
plate in 3–4 wk, aerial hyphae nearly lacking at the margin,
but dense in central areas, forming more or less radially
arranged spots; aged cultures only pale yellowish brown;
diffusing pigment lacking.
Habitat and host range: dead corticated twigs and
branches of Betula spp.
Distribution: common in North America (USA, Canada);
rare in Europe (France). The French collection investigated
in the present study is the first verified record from Europe.
Wehmeyer (1941) mentioned a European collection deposited in PC without providing further details, but considered
this record to be dubious due to the similarity of M. decorahensis to M. spodiaea, where ascospore appendages may
not be seen in old material.
Typification: USA, Iowa, Decorah, on corticated
branches of Betula sp., Aug. 1882, E. W. D. Holway (NY
922009, lectotype of Melanconis decorahensis here designated; NY 922008, isotype).
Additional selected specimens examined: FRANCE,
Ariège, Rimont, on twigs of Betula pendula, 14 July 2010,
J. Fournier, J.F. 10095, strains MD (from teleomorph), MED
(from anamorph) (WU 31808, culture CBS 131698). USA,
Alabama, Marion Co., Hamilton, Buttahatchee
(“Ballahatchie”) River, on twigs of Betula sp., 14 July
1925, L. E. Wehmeyer 3644 (DAOM 121211, NY). Iowa,
Decorah, on twigs of Betula sp., E. W. D. Holway, Ellis &
Everhart, N. Amer. Fungi. Ser. II: 1562 (NY, 3 specimens).
Massachusetts, Waverly, on twigs of Betula sp., 20 Sep.
1925, L. E. Wehmeyer 3806 (DAOM 121219, culture
CBS 159.26). Indiana, Kankakee River & Route # 43, on
twigs of Betula lutea, 3 Sep. 1938, L. E. Wehmeyer 3876
(DAOM 121187).
Notes: Melanconiella decorahensis is characterised by
dark brown ascospores lacking appendages, pip-shaped dark
brown conidia and growth on Betula spp. It is most similar
to M. spodiaea, which mainly differs by the presence of caplike ascospore appendages, differently shaped conidia and
different hosts (Carpinus spp.). Melanconis stilbostoma,
which is common on Betula spp., mainly differs by hyaline
ascospores and whitish ectostromatic discs. The anamorphteleomorph connection was proven by culture (Wehmeyer
1926) and DNA data.
We have not been able to study the type of M. decorahensis var. subviridis (0M. subviridis), but according to
Wehmeyer (1941) it represents merely a well-developed
specimen of M. decorahensis.
Fungal Diversity (2012) 57:1–44
Melanconiella echinata Voglmayr & Jaklitsch, sp. nov.
(Fig. 9)
MycoBank MB 800117
Etymology: referring to the ostioles projecting from the
prominently erumpent ectostromatic discs, resulting in a
spiny appearance.
Pseudostromata 1.2–2.5 mm diam, projecting 0.3–0.6(–
0.8) mm, well-defined, distinct, circular, causing a ring-like
elevation of the bark surrounding the ectostromatic disc;
perithecial bumps indistinct. Ectostromatic disc 0.4–0.9(–
1.3) mm diam, well-developed, distinctly projecting, often
pulvinate, with circular, angular or ellipsoid outline, cream,
yellow, honey, cinnamon, orange brown to brown. Central
column bright olive-yellow when young, later olive-green to
olive-brown. Entostroma well-developed, compact, of compacted hyphae, olive-green or olive-brown. Ostioles 1–25(–
30) per disc, 90–190 μm diam, typically regularly arranged
in the disc, cream to brown with shiny black tips, distinctly
projecting, papillate or conical, tips flat or rounded.
Perithecia 0.4–0.8 mm diam. Asci fusoid, (66–)72–81(–
83)×(13–)14–16.5(–18) μm (n033), containing 8 uni- or
biseriate ascospores, with a distinct apical ring, ca. 3 μm
diam, 2 μm high. Ascospores hyaline, fusoid, slightly
curved, slightly to distinctly constricted at the septum,
(18–)20–22(–26)×(6.0–)6.5–7.5(–8.7) μm, l/w0(2.5–)2.8–
3.2(–3.7) (n0121); ends subacute to narrowly rounded, with
hyaline cap-like appendages 1–1.5 μm long, 2–2.5 μm
wide, soon fading in microscope mounts; cells slightly
dimorphic, with four large guttules when fresh, mostly disappearing in herbarium specimens; wall ca. 0.4 μm thick,
sometimes distinctly swelling at the septum.
Anamorph: not observed.
Habitat and host range: dead corticated twigs and
branches of Carpinus caroliniana; probably rare.
Distribution: North America (USA).
Type: USA, New York, Ringwood, Lloyd-Cornell
Preserve, on corticated branches of Carpinus caroliniana,
6 Sep. 1952, R. M. Horner (DAOM 121196, holotype).
Additional specimens examined: USA, Pennsylvania,
West Chester, on Carpinus caroliniana, Aug. 1883, Haines
& Everhart, issued as Melanconis chrysostroma in Ellis &
Everhart, N. Amer. Fungi. Ser. II: 1563 (GZU).
Notes: As no recent collections have been available, only
the ITS-LSU region could be obtained for M. echinata, and
it could not be included in the combined phylogenetic
analyses. The LSU demonstrates that the fungus belongs
to Melanconiella, and that it is also distinct from all other
species (Fig. 1). Morphologically it fits well in
Melanconiella, except for its distinctly pustulate stromata,
which have not been observed in any other species of the
genus. Although otherwise being similar to M. elegans, M.
echinata is distinct due to its differently shaped, larger
Fungal Diversity (2012) 57:1–44
23
a
b
c
f
e
i
d
n
w
k
j
o
p
x
q
y
g
r
z
h
l
m
s
t
u
v
a1
b1
c1
d1
Fig. 9 Melanconiella echinata. a. Panoramic view of pseudostromata.
b. Pseudostroma in transverse section. c. Groups of detached perithecia
in well developed entostroma from below. d. Three pseudostromata in
surface view. e. Vertical section of pseudostroma. f. Ectostromatic disc
with projecting ostioles in side view. g–i. Young ectostromatic discs
and ostioles in surface view. j–l. Fully developed ectostromatic discs
and ostioles in surface view. m. Transverse section below ectostromatic
disc. n. Ascus. o. Ascus tip with apical ring. p–d1. Ascospores (p–s, z
showing appendages). n–t in water, u–d1 in 3 % KOH. a02 mm, b, c,
e–m0500 μm, d01 mm. Sources: a, b, d–y. DAOM 121196; c, z–d1.
Ellis & Everhart, N. Amer. Fungi. Ser. II: 1563 (GZU)
ascospores, its prominent erumpent ectostromatic discs
causing a coarsely pustulate bark surface similar to
Melanconis and its distinctly projecting ostioles. In addition,
the entostroma is well developed and of distinctly olive-
yellow to olive-green colour. Although the label of Ellis and
Everhart, N. Amer. Fungi 1563 states that the specimens
show no asci, fully mature perithecia were present in the
specimen from GZU, having asci and ascospores fully
24
in agreement with the type. Interestingly, this exsiccatum was issued as Melanconis chrysostroma, and it was
distributed in the same fascicle as the type collection of
Melanconiella ellisii (N. Amer. Fungi 1567, issued as
Valsa (Diaporthe) ellisii), which was collected at the
same locality. This suggests that Ellis and Everhart
considered these collections to represent distinct taxa.
Ellis and Everhart (1892) based their description of
Melanconis chrysostroma on the exsiccatum N. Amer.
Fungi 1563, applying the name with some doubts. In
the broad species concept of Wehmeyer (1941), the
specimen was listed under Melanconis chrysostroma
var. ellisii. We found no anamorph on the two collections examined.
Melanconiella elegans Voglmayr & Jaklitsch, sp. nov.
(Fig. 10)
MycoBank MB 800118
Etymology: referring to the regular yellowish ectostromatic discs with evenly distributed ostioles, providing an
elegant appearance.
Pseudostromata 1.2–3 mm diam, projecting to ca. 0.6(–
0.8) mm, well-defined, distinct, circular, or confluent with
densely disposed discs, causing small bumps in the bark;
perithecial bumps often distinct. Ectostromatic disc 0.2–
0.8 mm diam, rarely larger (1.6 mm), well-developed, distinctly projecting, often pulvinate, with typically circular or
angular, sometimes ellipsoid, fusoid or oblong outline,
pale or bright yellow to light brown, or concealed by
ostioles. Central column yellow, olive, grey or brownish. Entostroma well-developed, compact, of compacted
hyphae, dull yellow, olive or brownish. Ostioles 1–20
per disc, 30–140(–160) μm diam, typically evenly
spaced in the disc, brown with shiny black tips, slightly
projecting, papillate or conical, tips flat, rounded or
nearly acute. Perithecia 0.2–0.6 mm diam. Asci fusoid,
(49–)54–69(–77)×(9–)10–13(–15) μm (n067), containing 8 uni- or biseriate ascospores, with distinct apical
ring, 2.4–2.8 μm diam, 1.8–2.6 μm high. Ascospores
hyaline, fusoid, slightly to distinctly constricted at the
septum, (11.8–)14.5–17.0(–19.5) × (3.5–)4.3–5.3(–6.7)
μm, l/w 0(2.3–)2.9–3.7(–5.0) (n 0261); ends narrowly
rounded to subacute, appendages mostly indistinct; cells
slightly dimorphic, each containing 2 large guttules;
wall not swelling, ca. 0.4 μm thick.
Anamorph: discosporina-like. Conidiomata inconspicuous, typically only the central or eccentric ectostromatic disc
visible at the substrate surface or their presence indicated by
discharged conidial masses, 1–2.5 mm diam. Conidiogenous
cells phialidic, 10–25×2–3 μm, hyaline. Conidia hyaline,
fusoid or suballantoid, (7.5–)8.8–12.0(–14.3)×(2.2–)2.5–3.0
(–3.5) μm, l/w0(2.3–)2.9–4.4(–5.8) (n0215), multiguttulate
Fungal Diversity (2012) 57:1–44
with numerous small guttules; wall and hyaline gelatinous
sheath each ca. 0.3 μm thick.
Culture: Colony on MEA after 14 d at room temperature
ca. 14–35 mm diam, indistinctly zonate, surface slightly
farinose due to loosely disposed, small collapsed flocks of
aerial hyphae, first whitish, turning reddish-brown or rosy
mixed with yellow tones from the centre; after ca. 3 wk
purple pigment diffusing into the agar, colony turning dark
reddish-brown and eventually dark brown, grey to black;
diffusing pigment transient, unstable, grading off into
brown.
Habitat and host range: dead corticated twigs and
branches of Carpinus caroliniana; widespread.
Distribution: North America (USA).
Type: USA, Maryland, Prince George’s County,
Beltsville/Greenbelt, Beaver Dam Road, on corticated
branches of Carpinus caroliniana, 14 May 2003, W.
Jaklitsch & H. Voglmayr W.J. 2118, strain A.R. 3993 (BPI
843574, holotype; WU 31809, isotype; ex-holotype culture
CBS 131617), ex-holotype sequences JQ926265 (ITSLSU), JQ926336 (rpb2), JQ926402 (tef1).
Additional selected specimens examined (all on Carpinus
caroliniana): CANADA, Ontario, London, July 1891, J.
Dearness, Ellis & Everhart, N. Amer. Fungi. Ser. II: 2742
(K(M) 171590, issued as Diaporthe decipiens). USA,
M ar y l a n d , B e l t sv i l le , L i t t l e P a i nt B r a n c h , s o c .
Discosporina carpinicola, 20 May 1950, F. Petrak; Poelt
& Scheuer, Reliqu. Petrak. 2815 (GZU, issued as
Discosporina carpinicola). New York, vic. Ithaca, Arnot
Forest, 10 July 2002, L. Vasilyeva, strain A.R. 3830 (BPI
870990, culture CBS 131494). Tennessee, Gatlinburg, Great
Smoky Mountains National Park, Old Sugarlands Trail, 24
May 2003, W. Jaklitsch & H. Voglmayr, W.J. 2213 (BPI
872067). Virginia, Edinburg, George Washington National
Forest, Columbia Furnace/Wolf Gap, 27 May 2003, W.
Jaklitsch & H. Voglmayr, W.J. 2238 (WU 31810).
Notes: Asci, ascospores and anamorph of Melanconiella
elegans are similar to those of M. ellisii. Melanconiella
elegans has beautiful cream, yellowish to bright yellow
ectostromatic discs, which are uniform in shape and characterised by evenly scattered ostioles. The colour of its central
stromatic column is remarkably variable, ranging from
cream, light to dark grey, yellow, olive to brownish.
Melanconiella elegans is morphologically most similar to
the European M. carpinicola, which may be the closest
relative (Fig. 2). As the anamorph-teleomorph connection
could not be directly proven with pure cultures, DNA was
extracted from conidiomata present on the natural substrate.
Three independent DNA extractions from two specimens
(BPI 843574, BPI 872067) each confirmed the connection.
Morphological descriptions of Melanconis xanthostroma
by Wehmeyer (1937), later referred to as Melanconis chrysostroma var. ellisii by him (Wehmeyer 1941), are likely to
Fungal Diversity (2012) 57:1–44
25
a
e
b
g
f
m
l
h
u
v
a1
b1
d
i
o
n
t
c
c1
Fig. 10 Melanconiella elegans. a. Panoramic view of pseudostromata.
b, c. Pseudostromata in transverse section. d. Two pseudostromata in
surface view. e–i. Ectostromatic discs and ostioles in surface view. j, k.
Transverse section below ectostromatic disc. l. Ascus. m. Ascus tip
with apical ring. n–z. Ascospores. a1. Conidioma in surface view. b1.
j
p
w
k
q
r
x
s
y
z
d1
e1
f1
g1
h1
i1
j1
k1
l1
m1
Transverse section of conidioma, showing central column. c1. Conidiophores. d1–m1. Conidia. d1–f1, i1, j1 in water, all other mounts in
3 % KOH. a02 mm, b, c, a1, b10500 μm, d01 mm, e–k0300 μm, l–z,
c1–m1010 μm. Sources: a, b, g, h, k, b1. WU 31809; c, h1. BPI
872067; e, f, j, m–s. BPI 870990; i, l, t–a1, c1–g1, i1–m1. BPI 873574
26
be based on M. elegans. In his studies Wehmeyer (1937)
reported the production of large alpha conidia (16–30(–
37)×3.5–5 μm) in culture and similar conidia in a few
herbarium specimens. However, his cultures did not produce
the small beta-conidia he frequently observed on herbarium
specimens. In one of our collections of M. elegans (BPI
870990), numerous conidiomata containing such large conidia were present. However, these could not be confirmed
to represent a conidial state of M. elegans. A PDA culture
from ascospores of BPI 870990 produced conidia similar in
size (9–11×3–4 μm) to the small beta-conidia of Wehmeyer
(1937, 1941), which were confirmed to represent the anamorph of M. elegans in our study. Sequences obtained from
the large-spored anamorph on the natural substrate were
either impure (from two extraction trials containing conidia,
conidiophores and stromatic columns), or were revealed as
an unidentified, probably diaporthalean fungus not belonging to Melanconiella by a BLAST search (sequences from a
DNA extract obtained only from conidia). This indicates
that the conidia belong to a fungus that probably parasitizes
stromata of M. elegans.
The type collection of Discosporina carpinicola, which
was considered to be the anamorph of M. ellisii by Petrak
(1952b), includes the teleomorph of M. elegans. However,
morphologically the conidia present on the specimen are
more similar to those of M. ellisii. Considering common
co-occurrence and intermingled growth of teleomorphs and
anamorphs of different species on the same branches in
Melanconiella and the subtle morphological differences between the anamorphs of M. ellisii and M. elegans, it remains
uncertain to which species of Melanconiella Discosporina
carpinicola is connected. The species epithet is a later
homonym of Diaporthe carpinicola and therefore not available for use in Melanconiella.
Melanconiella ellisii (Rehm ex Ellis & Everh.) Voglmayr &
Jaklitsch, comb. nov. (Fig. 11)
MycoBank MB 800119
Basionym: Diaporthe ellisii Rehm ex Ellis & Everh., Bull
Torr Bot Club 10:89 (1883).
Synonyms: ?Discosporina carpinicola Petr., Sydowia
6:357 (1952).
Melanconis chrysostroma var. ellisii (Rehm ex Ellis &
Everh.) Wehm., Univ Michigan Stud Sci Ser 14:33 (1941).
Pseudostromata 1–2.6 mm diam, projecting up to ca.
0.7 mm, inconspicuous, circular, distinct or confluent; distinct perithecial bumps uncommon. Ectostromatic disc 0.2–
1.2(–2) mm diam, with circular, angular or oblong outline,
commonly surrounded by irregular edges or flaps of bark,
drab, light to dark grey or dark brown. Central column grey
or brownish. Entostroma typically indistinct, sometimes
compact above perithecia, of subhyaline hyphae. Ostioles
Fungal Diversity (2012) 57:1–44
1–15 per disc, 30–160(–170) μm diam at the base, typically
arranged at the disc margin, shiny black, plane, papillate or
conical and projecting up to 180 μm, tips flattened, rounded
or acute. Perithecia 0.2–0.5 mm diam. Asci fusoid, (39–)
50–69(–80)×(7–)9–11(–13) μm (n084), containing 8 unior biseriate ascospores, with distinct apical ring, 1.8–2.5 μm
diam, 1.4–2 μm high. Ascospores hyaline, fusoid, slightly to
distinctly constricted at the septum, (10–)12.5–16(–20)×
(3.5–)4.0–5.5(–6.5) μm, l/w0(2.2–)2.7–3.3(–4) (n0239);
ends narrowly rounded to subacute, mostly without appendages; cells mono- to slightly dimorphic, with (1–)2 large
and numerous small guttules; wall not swelling, ca. 0.3 μm
thick.
Anamorph: discosporina-like. Conidiomata inconspicuous, typically only their central or eccentric ectostromatic
disc visible on the substrate surface or their presence indicated by discharged conidial masses, 0.5–2 mm diam.
Conidiogenous cells phialidic, 10–18×2–3.5 μm, hyaline.
Conidia hyaline, fusoid, narrowly ellipsoid, elongate to
suballantoid, (8.5–)9.5–11.5(–13.0)× (2.5–)3.0–3.5(–4.0)
μm, l/w0(2.2–)2.8–3.9(–4.8) (n0148), multiguttulate, commonly with four to six large guttules; wall ca. 0.3 μm,
hyaline gelatinous sheath ca. 0.4 μm thick.
Habitat and host range: dead corticated twigs and
branches of Carpinus caroliniana; widespread.
Distribution: North America (USA, Canada).
Typification: USA, Pennsylvania, West Chester, on
Carpinus caroliniana, without date, Haines & Everhart,
Ellis & Everhart, N. Amer. Fungi. Ser. II: 1567 (M0177942, lectotype of Diaporthe ellisii here designated;
BPI bound, K(M) 171591, isotypes).
Additional selected specimens examined (all on Carpinus
caroliniana): USA, Maryland, Beltsville/Greenbelt, Beaver
Dam Road, 14 May 2003, W. Jaklitsch & H. Voglmayr, W.J.
2118a (BPI 883227; WU 31811). Maryland, Prince
George’s Co., Beltsville Agricultural Research Station,
Bottomland Research Forest, 3 June 2002, L. Vasilyeva
(BPI 843491). Tennessee, Great Smoky Mountains
National Park, vic. Cosby, 8 May 2006, L. Vasilyeva (BPI
878343).
Notes: Melanconiella ellisii, commonly referred to as
Melanconis chrysostroma var. ellisii, was previously considered to be a well-defined taxon in North America
(Wehmeyer 1941). However, the current study shows that
it is a heterogeneous taxon, and M. elegans is separated as a
distinct species. Melanconiella ellisii is characterised by its
drab, grey to dark brown ectostromatic discs of irregular
shape with ostioles inserted mostly at the margin of the
ectostromatic disc, which is reminiscent of the closely related M. spodiaea. Unfortunately, no pure cultures have been
available for M. ellisii, but DNA extraction from teleomorphs and anamorphs of three collections of M. ellisii
revealed consistent results.
Fungal Diversity (2012) 57:1–44
27
a
b
d
f
e
l
k
g
m
s
t
h
u
c1
Fig. 11 Melanconiella ellisii. a. Panoramic view of pseudostromata. b.
Pseudostroma in transverse section. c. Six pseudostromata in surface
view. d–i. Ectostromatic discs and ostioles in surface view. j. Transverse section below ectostromatic disc. k. Ascus. l. Ascus tip with
apical ring. m–z. Ascospores (m, r showing appendages). a1. Conidioma in surface view. b1. Transverse section of conidioma showing
i
o
n
a1
b1
c
p
w
v
j
q
x
r
y
z
d1
e1
f1
g1
h1
i1
j1
k1
l1
m1
central column. c1. Conidiophores. d1–m1. Conidia. All mounts in
3 % KOH. a02 mm, b, d–i0500 μm, c, j01 mm, k–z, c1–m1010 μm,
a1, b10300 μm. Sources: a, d, e, l, s–v. BPI 843491; b, f, j, k, w–z. BPI
878343; c–g, m, a1–b1, i1–m1. M-0177942 (lectotype); h, i, n–r. K(M)
171591 (isotype); c1–h1. BPI 883227)
28
Ellis and Everhart (1892) described the conidia as fournucleate, measuring 8–11× 2–2.5 μm. On the original collection deposited in Kew (K(M) 171591), the conidia are
distinctly wider (4–5.5 μm) than those of the collection
deposited in Munich (M-0177942), as well as in our collection BPI 883227 (WJ 2118a) for which the connection was
confirmed by sequence data. It remains to be verified whether the conidia of M. ellisii show a variable width or whether
these wider conidia belong to a different taxon.
For reasons of priority, Petrak (1952b) proposed that the
species epithet carpinigera should be used for the species.
However, this is a misapplied name, as the type specimen of
the basionym, Diatrype carpinigera, is clearly not a diaporthalean fungus but rather a Valsaria (see below). He also
described Discosporina carpinicola as the anamorph of M.
ellisii, but the type specimen contains the teleomorph of
only M. elegans. As D. carpinicola is morphologically more
similar to the anamorph of M. ellisii, it is here considered a
synonym of the latter.
The recent record of M. ellisii (as Melanconis carpinigera) from Carpinus cordata near Vladivostok, Russian Far
East by Vasilyeva and Stephenson (2010) cannot be evaluated without detailed study of the specimen.
Melanconiella flavovirens (G.H. Otth) Voglmayr &
Jaklitsch, comb. nov. (Fig. 12)
MycoBank MB 800120
Basionym: Diaporthe flavovirens G.H. Otth, Mitt naturf
Ges Bern 1868:47 (1869).
Synonyms: Melanconis flavovirens (G.H. Otth) Wehm.,
Mycologia 29:602 (1937).
?Diaporthe flavovirens var. tetraspora G.H. Otth, Mitt
naturf Ges Bern 1868:47 (1869).
Diaporthe olivaestroma (Cooke) Berl. & Voglino [as
‘elaeostroma’], in Saccardo, Syll Fung, Addit I–IV:105
(1886).
Diaporthe sulphurea Fuckel, Jb Nassau Ver Naturk 23–
24:205 (1870) [1869–70] (type not seen).
Discodiaporthe sulphurea (Fuckel) Petr., Hedwigia
62:294 (1921).
Discosporina sulphurea (Sacc.) Petr., Sydowia 16:187
(1963) [1962].
Discosporium sulphureum (Sacc.) Petr., Hedwigia 62:
292 (1921).
Melanconis sulphurea (Fuckel) Petr., Ann Mycol 21:321
(1923).
Myxosporium sulphureum Sacc., Syll Fung 3:725 (1884).
Valsa olivaestroma Cooke, Grevillea 14(no. 70):48
(1885).
Pseudostromata 1.5–3.4 mm diam, mostly irregularly
confluent, with slightly projecting centre, sometimes welldefined, circular with circinately arranged perithecial
Fungal Diversity (2012) 57:1–44
bumps. Ectostromatic disc 0.3–1.3 mm diam, circular, elliptic or oblong, often pulvinate, in this case not surrounded by
bark, powdery, mostly dull yellowish green to olive, but also
cream or yellow-brown, eventually black when old. Central
column grey, olive-brown, dull green or lacking. Entostroma
comprising olive or yellow mycelium, more or less well
developed. Ostioles 1–15(–25) per disc, emerging in various
positions in the disc, 40–160 μm diam, shiny black, slightly
papillate, less commonly plane, sometimes cylindrical and
projecting up to 150 μm when the disc disintegrates.
Perithecia 0.3–0.6(–0.8) mm diam. Asci fusoid, (84–)95–
107(–114) × (13–)14–17(–18) μm (n 031), containing
8 biseriate ascospores, with distinct apical ring, 3.2–
4.0 μm diam, 2.5–3.3 μm high. Ascospores hyaline, inequilaterally ellipsoid or broadly fusoid, not to slightly constricted at the septum, (17–)20.5–24.5(–30)×(7.0–)7.5–8.8
(–9.8) μm, l/w0(2.1–)2.5–3.0(–3.7) (n0134); ends narrowly rounded or subacute, with distinct persistent, knob-like
hyaline appendages 0.7–1.4 μm long, 1.6–2.4 μm wide;
cells mostly monomorphic, distinctly triangular-ovate in
outline, with one large and numerous small guttules;
wall not swelling, ca. 0.5–0.6 μm thick, at the septum
1.5–2 μm.
Anamorph: discosporina-like. Conidiomata 1.5–2.5 mm
diam, indistinct, mostly with central or eccentric stromatic
column; usually no dicharged conidial masses evident.
Conidiogenous cells phialidic, 16–27×2.5–3.5 μm, hyaline. Conidia hyaline, variable in shape, pip-shaped, narrowly ellipsoid, elongate to suballantoid, sometimes with a
distinct apiculus, (11–)12–15(–17.5)×(4.7–)5.0–5.5(–6.5)
μm, l/w0(1.9–)2.3–2.8(–3.3) (n065), multiguttulate, often
with one or two larger and numerous small guttules; wall
and hyaline gelatinous sheath each ca. 0.5 μm thick.
Culture: Colony on MEA after 17 d at room temperature
ca. 23 mm diam, after reaching a diam of 2–2.5 cm growth
discontinued or at most small irregular lobes formed; colony
dull greenish-yellow to yellow-brown with whitish margin,
turning dark brown to black from the centre; aerial hyphae
scant or abundant, causing a floccose surface; after 3–4 wk
pale to medium brown pigment diffusing into the agar;
odour reminiscent of fermenting yeast.
Habitat and host range: dead corticated twigs and
branches of Corylus avellana lying on the ground.
Distribution: Europe, probably also North America
(Canada, USA); common. Collections from North
American Corylus species need to be thoroughly studied
to evaluate whether they are conspecific.
Typification: SWITZERLAND, near Steffisburg, without
date, G. Otth 31 (B 700014743, lectotype of Diaporthe
flavovirens here designated; B 700014742, isotype).
SWITZERLAND, 2 May 1867, P. Morthier, Herb. Fuckel
56 (G 00111561, holotype of Diaporthe sulphurea).
SWITZERLAND, Jura, 31 March 1871, P. Morthier, Herb.
Fungal Diversity (2012) 57:1–44
29
a
b
d
x
f
e
j
i
c
q
y
g
h
k
l
m
n
o
p
r
s
t
u
v
w
z
a1
b1
c1
f1
g1
h1
i1
d1
j1
e1
k1
Fig. 12 Melanconiella flavovirens. a. Panoramic view of pseudostromata. b. Pseudostroma in transverse section. c. Two pseudostromata in
surface view. d–g. Ectostromatic discs and ostioles in surface view. h.
Transverse section below ectostromatic disc. i. Ascus. j. Ascus tip with
apical ring. k–w. Ascospores. x. Transverse section of conidioma. y.
Conidiophores. z–k1. Conidia. All mounts in water. a02 mm, b, c0
1 mm, d–h, x0500 μm, i–w, y–k1010 μm. Sources: a–d, j–n, x–k1.
WU 31822; e. WU 31815; f–i, o–w. WU 31816
Fuckel 27 (G 00111404, lectotype of Myxosporium sulphureum here designated). UK, England, Middlesex,
Hendon, M.C. Cooke 209 (K(M) 174473, holotype of
Valsa olivaestroma)
30
Additional specimens examined (all on Corylus avellana): AUSTRIA, Kärnten, St. Margareten im Rosental,
Sabosach, map grid 9452/3, 30 Oct. 1994, W. Jaklitsch,
W.J. 322 (WU 31812). ibid., shrubs between the village
and Stariwald, map grid 9452/4, 10 June 1993, W. Jaklitsch
(WU 15584). ibid., 27 Aug. 1993, W. Jaklitsch (WU 15510).
ibid., 19 Feb. 1994, W. Jaklitsch (WU 15376). ibid.,
Trieblach, below Kucher, map grid 9452/2, 25 March 1995,
W. Jaklitsch, W.J. 536 (WU 31813). ibid., Schwarzgupf, map
grid 9452/4, 10 Sep. 1998, W. Jaklitsch, W.J. 1202 (WU
31814). ibid., Wograda, map grid 9452/3, 27 Oct. 2000, W.
Jaklitsch, W.J. 1666 (WU 31815). ibid., Zabrde, map grid
9452/4, 17 Oct. 2011, H. Voglmayr, strain MFV2 (from teleomorph) (WU 31816). Niederösterreich, Orth an der Donau,
Tierboden, map grid 7866/3, 1 Nov. 2011, H. Voglmayr & W.
Jaklitsch, strain MFV4 (from teleomorph) (WU 31817).
Stockerau, Unterzögersdorf, map grid 7663/1, 24 April
2010, H. Voglmayr (WU 31818). Oberösterreich, Raab,
Rotes Kreuz, map grid 7648/1, 6 Sep. 2009, H. Voglmayr,
strain MFV (from teleomorph) (WU 31819, culture CBS
125598). St. Willibald, Aichet, map grid 7648/1, 3 June
2011, H. Voglmayr, strain MFV1 (from teleomorph)
(WU 31820). Steiermark, Pöllau, Schönauklamm, map
grid 8760/2, 12 Sep. 2002, W. Jaklitsch, W.J. 1952
(WU 31821). GERMANY, Reichartshausen, spring,
without date, L. Fuckel, Fungi Rhen. Exs. 2539 (K(M)
174474, issued as Diaporthe sulphurea). ITALY,
Lombardia, Vobarno, 21 Oct. 2011, H. Voglmayr & W.
Jaklitsch, strains MFV3 (from teleomorph), MFA (from
anamorph) (WU 31822). SWITZERLAND, Neuchâtel,
14 April 1869, P. Morthier, Herb. Barbey-Boissier 105
(BPI 617275 as Diaporthe sulphurea). same place, date
and collector (G 00111562 as Diaporthe sulphurea).
Notes: Melanconiella flavovirens is characterised by
growth on Corylus, by mostly large ectostromatic discs with
distinctly yellowish green tints and by broad ascospores of
cells that are distinctly triangular to ovate in outline and by
small persistent knob-like appendages. The anamorph is less
prominent than in other Melanconiella species, inconspicuous and rarely seen. Despite thorough studies of
several fresh collections, conidiomata were only found
in a single collection from Italy. The anamorph,
Myxosporium sulphureum, was formally described by
Saccardo (1884), who referred to the description by
Fuckel (1871). Apart from a few omissions, the wording
of the diagnosis (including measurements) in Saccardo
(1884) is identical with Fuckel’s description (1871:318–
319). Therefore, Myxosporium sulphureum should be
based on a collection studied by Fuckel, and the specimen from the Fuckel herbarium housed in G is here
designated as lectotype. The measurements of Fuckel
(1871), repeated in Saccardo (1884), agree with ours.
The anamorph-teleomorph connection was proven in the
Fungal Diversity (2012) 57:1–44
present study by DNA data from conidial and ascospore
cultures.
We examined two specimens of the type collection of
Diaporthe flavovirens preserved in B. Considering the
preservation and abundance of the collection, B
700014743 is here designated as lectotype. According
to the detailed description and the host (Fuckel 1870),
Diaporthe sulphurea is a synonym of Melanconiella
flavovirens, which has been confirmed by examination
of the type. Otth (1869) described a four-spored variety
of M. flavovirens, but no such specimen has been seen.
Melanconiella hyperopta (Nitschke ex G.H. Otth) Voglmayr
& Jaklitsch, comb. nov. (Fig. 13)
MycoBank MB 800121
Basionym: Diaporthe hyperopta Nitschke ex G.H. Otth,
Mitt naturf Ges Bern 1868:47 (1869).
Synonyms: Diaporthe decipiens Sacc, Fung Ven 4:6
(1875) (type not seen).
Melanconis hyperopta (Nitschke ex G.H. Otth) Wehm.,
Univ Michigan Stud Sci Ser 9:254 (1933).
Pseudostromata typically distinct, well-defined, with circular outline, 1.5–3.4 mm diam, projecting up to 0.5 mm;
perithecial bumps often evident; sometimes confluent and
irregular. Ectostromatic disc concave, flat or slightly pustulate, surrounded by bark or not, little projecting above the
perithecial level, 0.3–0.9 mm long, circular, fusoid to linear,
whitish, cream, pale or dull yellow to brown. Central column pale, yellowish, pale greyish brown or brownish.
Entostroma inconspicuous, pale, slightly lighter than bark
interior, comprising subhyaline to yellowish hyphae.
Ostioles 1–15 per disc, 60–160 μm diam, marginal or emergent within the disc, plane, discoid or papillate with rounded
or flattened apices. Perithecia 0.3–0.8 mm diam. Asci
fusoid, (73–)80–98(–111)×(12–)13–17(–21) μm (n071),
containing 8 biseriate ascospores, with distinct apical ring,
2.6–3.1 μm diam, 2.4–3 μm high. Ascospores hyaline,
fusoid, distinctly constricted at the septum, (17.7–)21–24(–
29.5)×(4.5–)5.5–6.8(–8.3) μm, l/w0(2.9–)3.3–4(–5.1) (n0
472); ends subacute to narrowly rounded, with hyaline caplike appendages 1.3–2.4 μm long, 3.5–5.6 μm wide, soon
fading in microscope mounts; cells dimorphic, upper cell
mostly larger, multiguttulate; wall not swelling, ca. 0.4 μm
thick.
Anamorph: discosporina-like. Conidiomata 1–1.5 mm
diam, often lined by a distinct, narrow blackish marginal
zone, with a central or eccentric stromatic column; at maturity covered by whitish discharged conidial masses.
Conidiogenous cells phialidic, 10–30×1.5–3 μm, hyaline.
Conidia hyaline, variable in shape, narrowly ellipsoid, elongate to slightly allantoid, (10–)11.5–14(–16.5)×(3.0–)3.8–
4.5(–5.0) μm, l/w 0(2.2–)2.7–3.5(–4.6) (n 0172),
Fungal Diversity (2012) 57:1–44
31
a
b
d
i
e
f
k
l
m
n
q
r
s
t
u
y
c1
d1
h
g
j
x
b1
c
o
v
z
e1
f1
p
w
a1
g1
Fig. 13 Melanconiella hyperopta. a. Panoramic view of pseudostromata. b. Pseudostroma in transverse section. c. Two pseudostromata in
surface view. d–g. Ectostromatic discs and ostioles in surface view. h.
Transverse section below ectostromatic disc. i. Ascus. j. Ascus tip with
apical ring. k–w. Ascospores. x. Conidioma in surface view. y. Trans-
h1
i1
j1
k1
l1
verse section of conidioma. z, a1. Conidiophores. b1–l1. Conidia. k,
l in 3 % KOH, all other mounts in water. a02 mm, b–h, x, y0500 μm,
i–w, z–l1010 μm. Sources: a. WU 31828; b–e. WU 31831; f, g, k, l.
Otth 115 (B, holotype); h, v. WU 31827; i, j, u, x–l1. WU 31836; m, n.
WU 31823; o–p. WU 31832; q–t. WU 31826; w. WU 31825
32
multiguttulate, often with one to four larger and few to
numerous small guttules; wall ca. 0.4 μm thick; sometimes
with an indistinct hyaline gelatinous sheath ca. 0.5 μm
thick.
Culture: Colony on MEA after 17 d at room temperature
ca. 75 mm diam, after ca. 20 d plate nearly entirely covered
(no further growth); colony not zonate, surface with distinct
radial to substellate arrangement of surface hyphae, aerial
hyphae irregularly arranged, inconspicuous or causing minute floccules on the surface; colony first white, turning
bright yellow to orange from the centre, eventually dull
orange; diffusing pigment lacking, agar slightly yellowish.
Habitat and host range: dead corticated twigs and
branches of Carpinus betulus mostly lying on the ground.
Distribution: Europe; common.
Typification: SWITZERLAND, near Bern, on corticated
branch of Carpinus betulus, without date, G. Otth 115 (B
700009156; lectotype designated by Wehmeyer (1933)).
Niederösterreich, Gaaden, Badener Bürgerspitalswald, map
grid 7963/1, on corticated branch of Carpinus betulus, 3
Dec. 2012, H. Voglmayr & I. Greilhuber, strain MHG (from
anamorph) (WU 31836, epitype here designated; ex-epitype culture CBS 132231), ex-epitype sequences JQ926285
(ITS-LSU), JQ926351 (rpb2), JQ926418 (tef1).
Additional specimens examined (all on Carpinus betulus): AUSTRIA, Burgenland, Neusiedl/See, Edelstal,
Spitzerberg, map grid 7967/2, 10 April 2010, H. Voglmayr
& I. Greilhuber, strain MCS (from teleomorph) (WU
31823). Kärnten, St. Margareten im Rosental, Trieblach,
near Ledra, map grid 9452/4, 1 May 2002, W. Jaklitsch,
W.J. 1878, strain A.R. 3832 (BPI 843521, culture CBS
131492). Niederösterreich, Mannersdorf am Leithagebirge,
Schweingraben, map grid 8065/2, soc. M. chrysodiscosporina (anamorph), 17 April 2010, H. Voglmayr & I.
Greilhuber (WU 31824). Mauerbach, near the cemetery,
map grid 7763/1, 28 Sep. 1996, W. Jaklitsch, W.J. 968
(WU 31825). ibid., 12 June 2011, W. Jaklitsch (WU
31826). Oberösterreich, Raab, Riedlhof, map grid 7647/2,
4 April 2010, H. Voglmayr, strain MCR (from teleomorph)
(WU 31827). Raab, Wetzlbach, map grid 7648/1, 4 April
2010, H. Voglmayr (WU 31828). St. Willibald, Loitzmayr,
map grid 7648/3, 8 April 2010, H. Voglmayr, strain MCL
(from teleomorph) (WU 31829, culture CBS 131697).
Wien, Döbling, Cobenzl, map grid 7763/2, 11 Feb. 2001,
W. Jaklitsch, W.J. 1715, strain A.R. 3619 (BPI 840939, WU
31830, culture CBS 131493). Döbling, Hermannskogel,
map grid 7763/2, soc. M. chrysodiscosporina (anamorph),
18 April 2010, H. Voglmayr, strain MCHE (from teleomorph) (WU 31831, culture CBS 131696). Landstraße,
Botanical Garden, map grid 7864/1, 31 March 2010, H.
Voglmayr, strain MCHBV (from teleomorph) (WU
31832). Liesing, Maurer Wald, map grid 7863/1, 3 Oct.
1998, W. Jaklitsch, W.J. 1224 (WU 31833). ibid., 22 April
Fungal Diversity (2012) 57:1–44
2006, W. Jaklitsch, W.J. 2911 (WU 31834). Ottakring,
Wilhelminenberg, Kreuzeichenwiese, map grid 7763/4, 19
March 2006, H. Voglmayr (WU 31835). GERMANY,
Bayern, München, Allach, Lochholz, 16 March 1998, L.
Beenken, Triebel, Microf. Exs. 273 (UPS 173895).
Sachsen-Anhalt, Eisleben, Oberrissdorfer Tal, May 1875,
J. Kunze, Fungi Sel. Exs. 122 (M-0177941, issued as
Diaporthe decipiens). SWEDEN, Skåne, Helsingborg,
Gyhults gård, 20 Jan. 1997, S.-Å. Hanson 25634 (UPS
200066). Skåne, Helsingborg, Frederiksdals
Friluftsmuseum, 28 Feb. 2012, S.-Å. Hanson 2012-007
(WU 31615). Skåne, Södra Mellby par, Stenshuvud, 20
Oct. 1990, K. & L. Holm 5607a (UPS 412043, UPS
412044, culture CBS 114600). UKRAINE, Stryj,
Podhorce, 27 Dec. 1916, F. Petrak, Fungi Polon. Exs. 637
(M-0157851, W).
Notes: Besides Melanconiella spodiaea, M. hyperopta is
the most common species of the genus occurring on
Carpinus betulus in Europe, where it is mainly found on
broken or fallen branches lying on the ground within forests.
It is commonly associated with M. chrysodiscosporina.
During attempts to establish anamorph-teleomorph connections in the present study, M. chrysodiscosporina was isolated several times from conidiomata closely associated with
stromata of M. hyperopta. Although the conidia of both
species are hyaline and of similar size, those of M. chrysodiscosporina can be distinguished by their more regular,
cylindrical shape and the presence of two large guttules in
the cell content. The anamorph-teleomorph connection was
proven by DNA data from conidial and ascospore cultures.
A culture of a Swedish collection (CBS 114600) had distinct
ITS-LSU, rpb2 and tef1 sequences (Fig. 2), but the microscopic features of the specimen (UPS 412044) agreed fully
with the other collections from Carpinus betulus. The collections from Carpinus orientalis are described as a new
variety based on slightly smaller ascospores and the different host (see below).
Wehmeyer (1933) proposed the combination Melanconis
hyperopta, but he included M. carpinicola in his species
concept. Finally, he synonymised M. hyperopta with M.
chrysostroma (Wehmeyer 1941). According to the spore
sizes given by Munk (1957) and Ellis and Ellis (1997), their
concept of Melanconis chrysostroma was evidently based
on collections of M. hyperopta. According to Gerhardt and
Hein (1979), the herbarium of Nitschke, containing duplicates of Otth’s material, is present in the herbarium in Berlin
(B). The spore size given in Otth (1869) is somewhat
smaller (18–20×4–5 μm) than that revealed for the authentic collection deposited in Berlin (20–26×5–7.5), which is
within the typical range of M. hyperopta. A similar case of
underestimated spore sizes given by Otth (1869) was also
recorded for Massaria pyri (Voglmayr and Jaklitsch 2011),
indicating that the measurements given by Otth may
Fungal Diversity (2012) 57:1–44
commonly be too small. To ensure nomenclatural stability, a
recent collection, for which cultures and sequences are
available, is designated here as epitype.
The basionym of M. hyperopta, Diaporthe hyperopta,
has been commonly ascribed to Nitschke, following
Otth (1869) who attributed the taxon to Nitschke.
However, there is no indication that Nitschke provided
a diagnosis, and Otth himself stated in the introduction
that the descriptions were intended to be provisional and
based on his own observations (Otth 1869:37–38).
Therefore, the author citation for D. hyperopta should
be Nitschke ex G.H. Otth.
Although no type specimen was available for study,
Diaporthe decipiens is clearly a synonym of M. hyperopta,
according to spore size and host. Diaporthe nigro-annulata
Kunze, considered a synonym of M. hyperopta by
Wehmeyer (1933), has apparently never been validly published; the schedae of the exsciccatum listed as type (Kunze,
Fungi Sel. Exs. 122) were issued as Diaporthe decipiens
Sacc., giving “Diap. nigro-annulata Johs. Kze. in sched.” as
a synonym. Re-examination of two specimens of Kunze,
Fungi Sel. Exs. 122 from Munich revealed one as M. hyperopta (M-0177941), whereas the other contains immature M.
carpinicola (M-0177940).
Melanconiella hyperopta var. orientalis Voglmayr &
Jaklitsch, var. nov. (Fig. 14)
MycoBank MB 800122
Etymology: referring to the epithet of the host, Carpinus
orientalis.
Pseudostromata 1.5–3 mm diam, typically distinct, welldefined, with circular outline, projecting up to 0.5 mm,
perithecial bumps often evident; sometimes confluent and
irregular. Ectostromatic disc typically flat, little projecting
above the perithecial level, 0.2–0.6(–0.8) mm long, circular
to linear, whitish, cream, light or pale yellow. Central column whitish, grey or brownish, entostroma inconspicuous.
Ostioles 1–12 per disc, 50–160 μm diam, marginal or disposed within the disc, plane, discoid or papillate with rounded or flattened apices. Perithecia 0.3–0.6 mm diam. Asci
fusoid, (84–)87–103(–107)×13–17(–18) μm (n012), containing 8 biseriate ascospores, with distinct apical ring, 2.5–
3.6 μm diam, 1.8–2.3 μm high. Ascospores hyaline, fusoid,
distinctly constricted at the septum, (17.5–)18.7–22(–24)×
(4.7–)5.7–6.7(–7.5) μm, l/w0(2.7–)3.0–3.5(–4.4) (n0110);
ends subacute to narrowly rounded, with hyaline cap-like
appendages 1–2 μm long, 2.7–4.2 μm wide, soon fading in
microscope mounts; cells dimorphic, upper cell mostly larger, multiguttulate; wall not swelling, ca. 0.4 μm thick.
Anamorph: discosporina-like. Conidiomata 0.9–1.5 mm
diam, visible as darker spots and lined by a darker marginal
zone, mostly with a central or eccentric stromatic column; at
33
maturity covered by whitish conidial masses. Conidiogenous
cells phialidic, 7–22×2–3.5 μm, hyaline. Conidia hyaline,
variable in shape, narrowly ellipsoid, elongate to slightly allantoid, (9.3–)10.5–12.7(–14.5)×(3.0–)3.5–4.0(–4.5)
μm, l/w0(2.2–)2.8–3.6(–4.4) (n071), multiguttulate, often with one to three larger and few to numerous small
guttules; wall and hyaline gelatinous sheath each ca.
0.5 μm thick.
Culture: Colony on MEA after 17 d at room temperature
83 mm diam, after 4–5 wk plate nearly entirely covered by
mycelium. Colony white, with pale brownish spots, or centre white, followed by a diffuse brownish, less commonly
rosy zone, otherwise not zonate, with distinct radial to substellate arrangement of surface hyphae; aerial hyphae scant,
sometimes abundant in the centre, diffusing pigment lacking; odour slightly musty.
Habitat and host range: dead corticated twigs and
branches of Carpinus orientalis; only found on branches
attached to the trees.
Distribution: South-eastern Europe, uncommon; only
known from Croatia but likely to co-occur elsewhere with
its host.
Type: CROATIA, Istria, Krnica, on corticated twigs of
Carpinus orientalis, 15 May 2010, H. Voglmayr & W.
Jaklitsch, strain MSK (from teleomorph) (WU 31837, holotype; ex-holotype culture CBS 131705), ex-holotype
sequences JQ926286 (ITS-LSU), JQ926354 (rpb2),
JQ926421 (tef1).
Additional specimen examined: CROATIA, Istria, Poreč,
Gedici, on corticated twigs of Carpinus orientalis, 1 Nov.
2010, H. Voglmayr & I. Greilhuber, strains MHP (from
teleomorph), MHVA (from anamorph) (WU 31838).
Notes: Melanconiella hyperopta var. orientalis is similar
to the closely related M. hyperopta, from which it differs
mainly by host, slightly smaller ascospores and conidia and
ITS, rpb2 and tef1 sequences. As these differences are
subtle, the fungus on Carpinus orientalis is described as a
new variety of M. hyperopta. The spores and conidia exhibit
pronounced drought resistance, as most were still vital after
storage for more than one year. The anamorph-teleomorph
connection was proven by DNA data from conidial and
ascospore cultures.
Melanconiella meridionalis Voglmayr & Jaklitsch, sp. nov.
(Fig. 15)
MycoBank MB 800123
Etymology: referring to the southern distribution within
Europe.
Pseudostromata 0.8–2.3 mm diam, projecting up to ca.
0.5 mm, inconspicuous, indistinct, confluent, less commonly distinct and circular, causing minute bumps in the bark;
perithecial bumps distinct or not. Ectostromatic disc
34
Fungal Diversity (2012) 57:1–44
a
b
d
j
a1
s
f
e
l
k
t
c
g
n
m
u
v
b1
Fig. 14 Melanconiella hyperopta var. orientalis. a. Panoramic view of
pseudostromata. b. Two pseudostromata in transverse section. c. Three
pseudostromata in surface view. d–h. Ectostromatic discs and ostioles
in surface view. i. Transverse section below ectostromatic disc. j.
Ascus. k. Ascus tip with apical ring. l–z. Ascospores. a1. Conidioma
o
w
c1
i
h
p
x
q
r
y
z
d1
e1
f1
g1
h1
i1
j1
k1
in surface view. b1. Transverse section of conidioma showing central
column. c1. Conidiophores. d1–k1. Conidia. All mounts in water. a0
2 mm, b01 mm, c0500 μm, d–i, a1, b10300 μm, j–z, c1–k1010 μm.
Sources: a, b, h, i, v–z. WU 31837; c–g, j–u, a1–k1. WU 31838
Fungal Diversity (2012) 57:1–44
35
a
b
d
c
e
f
h
g
j
i
k
l
m
n
o
p
q
u
r
v
w
x
s
y
z
Fig. 15 Melanconiella meridionalis. a. Panoramic view of pseudostromata. b. Pseudostroma in transverse section. c. Group of pseudostromata
in surface view. d–g. Ectostromatic discs and ostioles in surface view. h.
Transverse section below ectostromatic disc. i. Ascus. j. Ascus tip with
apical ring. k–q. Ascospores. r, s. Conidiomata in surface view. t.
t
a1
b1
Transverse section of conidioma. u. Conidiophores. v–b1. Conidia. All
mounts in water. a02 mm, b–h, r0500 μm, s, t0200 μm, i–q, u–b10
10 μm. Sources: a, c, d–f, r, s, u. WU 31845; b, g–j. WU 31844; k–p. WU
31849; q. WU 31846; t. WU 31839; v–b1. WU 31840
36
typically minute, 0.1–0.6 mm long, circular, oblong or linear, whitish, cream, pale yellowish to greyish or greyish
brown, or concealed by ostioles. Central column (pale) grey
or greyish brown. Entostroma whitish, of hyaline hyphae,
scant or well-developed. Ostioles few, 1–5 per disc, commonly only 1–3, rarely 7, 50–125(–160) μm diam, shiny
black, plane or slightly papillate with flattened or rounded
tips. Perithecia valsoid around the column or solitary, 0.2–
0.5(–0.8) mm diam. Asci broadly cylindrical to fusoid, (85–)
91–104(–115)×(13–)15–18(–21) μm (n031), containing
8 uni- or irregularly biseriate ascospores, with distinct apical
ring, 3.5–4.4 μm diam, 2.4–3 μm high. Ascospores hyaline,
fusoid, constricted at the septum, (20–)23–28(–38)×(5.0–)
6.5–8.0(–9.5) μm, l/w0(2.8–)3.2–3.9(–4.6) (n090); ends
narrowly rounded, with hyaline persistent cap-like appendages 1–1.8 μm long, 2.8–3.4 μm wide; cells dimorphic,
upper cell mostly larger, with (1–)2 large and numerous
small guttules; wall not swelling, 0.4 μm thick.
Anamorph: discosporina-like. Conidiomata 0.4–1.5 mm
diam, visible as dark brown spots lined by a darker marginal
zone, with a central or eccentric stromatic column; at maturity covered by whitish discharged conidial masses.
Conidiogenous cells phialidic, 19–37×3–4 μm, hyaline.
Conidia hyaline, cylindrical to suballantoid, (13–)15–17.5
(–19.5)×(5.8–)6.3–7.0(–8.0) μm, l/w0(1.7–)2.2–2.7(–3.2)
(n0120), with (1–)2 large and numerous small guttules;
wall ca. 0.4–0.6 μm thick, with prominent gelatinous sheath
0.8–1.2 μm thick.
Culture: On MEA growth conspicuously slow, colony
reaching a diam of ca. 16 mm after 17 d and ca. 22 mm
after 4 wk, white with diffuse yellow centre, dense, compact, thick; aerial hyphae short, forming delicate flocks;
diffusing pigment lacking; odour slightly yeast-like.
Habitat and host range: dead corticated twigs and
branches of Ostrya carpinifolia mostly still attached to the
tree.
Distribution: Southern Europe; known from Austria,
Croatia, Greece, Italy and Slovenia; common in the natural
distribution range of its host.
Type: AUSTRIA, Kärnten, St. Margareten im Rosental,
on corticated branches of Ostrya carpinifolia, map grid
9452/4, 12 May 2010, H. Voglmayr, strain MOM (from
teleomorph) (WU 31839, holotype; ex-holotype culture
CBS 131704), ex-holotype sequences JQ926291 (ITSLSU), JQ926357 (rpb2), JQ926424 (tef1).
Additional specimens examined (all on Ostrya carpinifolia): AUSTRIA, Kärnten, St. Margareten im Rosental,
Novina, map grid 9452/4, 3 July 2007, W. Jaklitsch, W.J.
3106 (WU 31842). ibid., shrubs W of Stariwald, map grid
9452/4, 17 May 2010, H. Voglmayr & W. Jaklitsch, strain
MOA (from anamorph) (WU 31840). ibid, map grid 9452/4,
10 Sep. 2010, W. Jaklitsch (WU 31841). ibid., Wograda,
map grid 9452/3, 13 Sep. 2001, W. Jaklitsch, W.J. 1810
Fungal Diversity (2012) 57:1–44
(WU 31843). CROATIA, Primorsko-goranska (Istria), nature park Učka, Veprinac WSW Opatija, 15 May 2010, H.
Voglmayr & W. Jaklitsch, strain MOU (from teleomorph)
(WU 31844). Vela Učka, Poklon, 15 May 2010, H.
Voglmayr & W. Jaklitsch, strain MOK (from teleomorph)
(WU 31845). GREECE, Kerkyra (Corfu), Kastellani near
Troumpettas, 21 April 2012, H. Voglmayr & W. Jaklitsch
(WU 32003). ITALY, Toscana, Grosseto, Sovana, 29 July
2009, H. Voglmayr, W. Jaklitsch & W. Gams, strain MOSO
(from teleomorph) (WU 31846). Trentino, Nembia, 19 Oct.
2011, H. Voglmayr & W. Jaklitsch (WU 31847).
SLOVENIA, Vipava, Mt. Nanos massif, Rebrnice NE
Lozice, 13 June 2009, H. Voglmayr & I. Greilhuber, strain
MON (from teleomorph) (WU 31848). Divača, Škocjan, 12
June 2009, H. Voglmayr & I. Greilhuber, strain MOS (from
teleomorph) (WU 31849). Divača, Škocjanska jama, 12
June 2009, H. Voglmayr & I. Greilhuber, strain MOSJ (from
teleomorph) (WU 31850, culture CBS 125599).
Notes: Melanconiella meridionalis is characterised by its
large, mostly biguttulate ascospores with dimorphic cells
and the growth on Ostrya carpinifolia. In addition, it has
the least developed stroma of all Melanconiella species. It is
closely related to the North American Melanconiella
ostryae, which differs by distinctly smaller ascospores
(15–23×4–8 μm), dark brown conidia with a lighter equatorial zone and the different host O. virginiana. The
anamorph-teleomorph connection was proven by DNA data
from conidial and ascospore cultures.
Melanconiella ostryae (Dearn.) Voglmayr & Jaklitsch,
comb. nov. (Fig. 16)
MycoBank MB 800124
Basionym: Diaporthe ostryae Dearn., Mycologia 18:246
(1926).
Synonym: Melanconis ostryae (Dearn.) Wehm.,
Mycologia 29:601 (1937).
Pseudostromata 1–2.2 mm diam, typically inconspicuous, sometimes distinct, circular, slightly projecting to ca.
0.5 mm, perithecial bumps uncommon. Ectostromatic disc
small, well-defined, with circular or elliptic outline, 0.1–
0.5 mm long, whitish, cream, grey to pale greyish brown.
Central column whitish, grey or brownish. Entostroma
whitish or cream, of hyaline hyphae. Ostioles 1–5(–7) per
disc, variously arranged, 40–125 μm diam, shiny black, flat
or papillate. Perithecia 0.2–0.5 mm diam. Asci cylindrical to
fusoid, (55–)77–100(–120)×10–13(–14) μm (n057), containing 8 irregularly biseriate ascospores, with apical ring
distinct when fresh, 2.8–3 μm diam, 2.2–3 μm high, only
faintly visible in older herbarium specimens. Ascospores
hyaline, fusoid, slightly to not constricted at the septum,
(14.5–)16.5–19(–23)×(4.0–)5.5–7.0(–8.3) μm, l/w0(2.1–)
2.5–3.3(–4.2) (n0209); ends narrowly rounded to subacute,
Fungal Diversity (2012) 57:1–44
37
a
b
d
f
e
k
j
i
c
r
s
m
l
t
g
n
v
u
h
o
p
q
w
x
y
c1
z
d1
a1
e1
f1
b1
g1
h1
i1
Fig. 16 Melanconiella ostryae. a. Panoramic view of pseudostromata.
b. Pseudostroma in transverse section. c. Three pseudostromata in
surface view. d–g. Ectostromatic discs and ostioles in surface view.
h. Transverse section below ectostromatic disc. i. Ascus. j. Ascus tip
with apical ring. k–y. Ascospores (k, u–y showing appendages). z, a1.
Conidiomata in surface view. b1. Transverse section of conidioma,
j1
k1
showing central column. c1, d1. Conidiophores. e1–k1. Conidia. k–v
in 3 % KOH, all other mounts in water. a02 mm, b, c, z–b10500 μm,
d–h0200 μm, i–y, c1–k1010 μm. Sources: a, e–q, b1. Bowser 37–102
(NY); b–d, u, v. Dearness 27 July 1893, Ellis 2162 (NY); r–t, a1.
M.E.B.B. 4365 (NY); w–y. Ellis 2162B (NY); z. TRTC 44844 (NY);
c1–k1. BPI 615036
38
with hyaline cap-like appendages 0.6–2 μm long, 2.2–
3.8 μm wide, soon disappearing in microscope mounts; cells
slightly dimorphic, upper cell mostly larger, multiguttulate,
sometimes with one large and numerous small guttules per
cell; wall not swelling, ca. 0.4 μm thick.
Anamorph: melanconium-like. Conidiomata 1–1.5 mm
diam, margins not visible on the surface, with central or
eccentric stromatic column; at maturity covered by black
discharged conidial masses. Conidiogenous cells annellidic,
20–32×3–5 μm, hyaline. Conidia dark brown, with a light
brown equatorial zone ca. 1.5–2.2 μm wide, ellipsoid,
slightly constricted in the middle, (12–)13–15.5(–17) ×
(6.2–)7.0–8.0(–8.8) μm, l/w0(1.6–)1.7–2.1(–2.4) (n076),
without guttules or granules; wall and hyaline gelatinous
sheath each ca. 0.5–0.7 μm thick.
Habitat and host range: dead corticated twigs and
branches of Ostrya virginiana.
Distribution: North America (USA, Canada); common.
Typification: CANADA, Ontario, Mt. Pleasant near
London, on corticated branches of Ostrya virginiana, 6
July 1904, J. Dearness 3152 (DAOM, lectotype of
Diaporthe ostryae here designated); same place, date and
collector, herbarium L. E. Wehmeyer 3278 (DAOM
120563, isotype).
Additional specimens examined (all on Ostrya virginiana): CANADA, Ontario, London, 27 July 1893, J.
Dearness, (NY, 2 specimens: Ellis 2162 & 2162B). ibid.,
without date, J. Dearness, Ellis & Everhart, N. Amer. Fungi.
Ser. II: 3430 (NY). Huron Distr., Bruce Co., Brinkman
Corner, 31 May 1964, R. F. Cain, TRTC 44844 (NY).
Bruce Co., Garrick twp., Con. XIII, Lot 31, 22 July 1960,
R. L. Bowser 37-102 (NY). York Co., Nashville, 2 May
1954, R. F. Cain (NY). USA, Vermont, Lamoille Co.,
Stowe, Covered Bridge Road, 2 July 1964, H. E. & M. E.
Bigelow M.E.B.B. 4191 (NY). ibid., Pinnacle Trail, 21 July
1964, H. E. & M. E. Bigelow M.E.B.B. 4365 (NY).
Notes: Melanconiella ostryae is characterised by growth
on Ostrya virginiana and dark brown conidia with a distinctly lighter brown equatorial zone, which is unique in
Melanconiella. Due to the dark brown conidia, which are
produced in large numbers, colonised twigs are blackened,
providing a strong contrast to the whitish ectostromatic
discs. In several collections, no black ostioles were visible
in the ectostromatic discs, and sections revealed that no or
only aborted perithecia were present. Due to the lack of
fresh collections, no cultures and sequences could be
obtained from the anamorph in the present study.
However, the anamorph-teleomorph connection was experimentally proven in pure culture studies by Wehmeyer
(1937). Melanconiella ostryae is closely related to M. meridionalis, which differs by distinctly larger ascospores (20–
38×5–10 μm), hyaline conidia, a different host (O. carpinifolia) and European distribution.
Fungal Diversity (2012) 57:1–44
Melanconiella spodiaea (Tul.) Sacc., Syll Fung 1:740
(1882). (Fig. 17)
MycoBank MB210704
Basionym: Melanconis spodiaea Tul., Ann Sci nat, ser 4,
5:109 (1856) (type not seen).
Pseudostromata inconspicuous, 1.3–3 mm diam, little
projecting, circular, perithecial contours typically not evident. Ectostromatic disc 0.25–1.1 mm long, mostly oblong,
sometimes circular or triangular, generally laterally enclosed
by bark, projecting 0.1–0.5 mm, often concealed by densely
disposed ostioles, light to dark grey, greyish brown, light to
dull or dark brown. Central column well-defined, light to
dark grey, greyish brown or grey-olive, perforated by
ostioles. Entostroma of subhyaline or yellowish hyphae.
Ostioles often in 1 or 2 rows, 1–15 per disc, 60–190 μm
diam, shiny black, stout, papillate to short cylindrical, with
rounded, sometimes flattened apices, distinct or fused in the
disc. Perithecia 0.3–0.5 mm diam, up to 20 per stroma. Asci
cylindrical, (104–)110–120(–121)× (14–)15–17(–18) μm
(n016), containing 8 uniseriate ascospores, with distinct
apical ring, 3–3.8 μm diam, 1–1.5 μm high. Ascospores
dark umber brown, broadly ellipsoid, slightly constricted
at the straight to slightly oblique septum, (13.5–)15.5–18.3
(–19.5)×(6.5–)7.5–8.8(–9.8) μm, l/w0(1.6–)1.9–2.3(–2.5)
(n082); ends broadly rounded, with broad, persistent, hyaline cap-like appendages 1–1.7 μm long, 4–5.5 μm wide;
cells monomorphic to slightly dimorphic, multiguttulate,
often with one large guttule almost entirely filling the lumen
and few peripheral small guttules; wall not swelling, ca.
0.7 μm thick.
Anamorph: melanconium-like. Conidiomata 1–4 mm
diam, blackish, mostly with central or eccentric stromatic
column; at maturity covered by black discharged conidial
masses. Conidiogenous cells annellidic, 20–29×2.5–5 μm,
hyaline. Conidia dark umber brown, variable in shape,
ovoid, obovoid, ellipsoid, oblong, sometimes shmooshaped or slightly constricted in the middle, (12.5–)13.3–
15.2(–16.5)×(7.0–)7.5–8.5(–9.0) μm, l/w0(1.4–)1.6–1.9(–
2.1) (n031), multiguttulate; wall and gelatinous sheath each
ca. 0.7 μm thick.
Culture: Colony on MEA after 17 d at room temperature
ca. 64 mm diam, with continuous or distinctly lobed margin,
surface whitish, farinose or granulose due to aerial hyphae
that form densely disposed floccules; floccules producing
hyaline, rosy to violaceous brown drops; reverse and agar
turning rosy, later reddish brown 9–10DE5–8 by a diffusing
pigment; eventually colony reverse dark red, purple to nearly black, surface dull whitish to grey; agar decolourised to
pale brownish; odour typically distinctly musty (like fermenting fruits).
Habitat and host range: dead corticated twigs and
branches of Carpinus betulus and C. orientalis.
Fungal Diversity (2012) 57:1–44
39
a
b
d
e
j
i
f
l
k
q
c
r
s
h
g
m
n
o
p
t
u
v
w
a1
x
b1
y
c1
d1
z
e1
f1
Fig. 17 Melanconiella spodiaea. a. Panoramic view of pseudostromata. b. Two pseudostromata in transverse section. c, h. Transverse
sections below ectostromatic disc. d–g. Ectostromatic discs and
ostioles in surface view. i. Ascus. j. Ascus tip with apical ring. k–w.
Ascospores. x, y. Conidiomata in surface view. z. Transverse section of
g1
h1
two conidiomata, showing central columns. a1. Conidiophores. b1–h1.
Conidia. All mounts in water. a, x02 mm, b–h, y, z0500 μm, i–w, a1–
h1010 μm. Sources: a–d, i, p–u, x–h1. WU 31854; e–h, v, w. WU
31855; j–o. WU 31614
40
Distribution: Europe; common and widespread.
Specimens examined: On Carpinus betulus: AUSTRIA,
Burgenland, Hornstein, Lebzelterberg, map grid 8064/4, 11
Sep. 2010, H. Voglmayr, strain MSH (from teleomorph)
(WU 31851). Kärnten, St. Margareten im Rosental, Ledra,
map grid 9452/4, 7 June 1996, W. Jaklitsch, W.J. 893 (WU
31852). Niederösterreich, Michelbach Markt, Mayerhöfen,
Hegerberg, map grid 7860/4, 20 June 2009, W. Jaklitsch &
H. Voglmayr, strain SPOD (WU 31853, culture CBS
125641). ibid., 3 June 2000, W. Jaklitsch, W.J. 1485, strain
A.R. 3462 (BPI 747944, culture CBS 109762). Mauerbach,
near the cemetery, map grid 7763/1, 12 Nov. 2011, W.
Jaklitsch, strain SPOD1 (from anamorph) (WU 31854).
ibid., 22 Aug. 1992, W. Jaklitsch (WU 16067). Hainbuch,
map grid 7763/1, 21 Aug. 1993, W. Jaklitsch (WU 15993).
Gaaden, Badener Bürgerspitalswald, map grid 7963/1, 3
Dec. 2012, H. Voglmayr & I. Greilhuber (WU 31614).
Oberösterreich, Raab, Großrothmayr, map grid 7647/2, 4
April 2010, H. Voglmayr (WU 31855). Wien, Döbling,
Hermannskogel, map grid 7763/2, 15. Feb. 1998, W.
Jaklitsch, W.J. 1135 (WU 31856). ibid., 27 March 1999,
W. Jaklitsch, W.J. 1295 (WU 31857). Hietzing, Lainzer
Tiergarten, map grid 7863/1, 28 June 1992, W. Jaklitsch
(WU 15204). Liesing, Maurerwald, map grid 7863/1, 10
July 1993, W. Jaklitsch (WU 15037). On Carpinus orientalis: CROATIA, Istria, Vrsar, soc. M. chrysorientalis, 14 May
Fungal Diversity (2012) 57:1–44
2010, H. Voglmayr & W. Jaklitsch, strain MVS (from anamorph) (WU 31858). GREECE, Kerkyra (Corfu), ESE Ano
Korakiana, W Ag. Vasilios, 23 April 2012, H. Voglmayr &
W. Jaklitsch (WU 32004).
Notes: Melanconiella spodiaea is the most common and
conspicuous species of the genus in Europe. It is often
associated with Prosthecium ellipsosporum, which appears
to be rarer. Melanconiella spodiaea is most similar to M.
decorahensis, which has dark brown ascospores of similar
size, but differs by the absence of ascospore appendages, by
pip-shaped conidia, an ectostroma of light to dark grey or
brown colour and different hosts (Betula spp.). Colonised
twigs are often distinctly blackened by numerous dark
brown conidia and ascospores. The anamorph-teleomorph
connection was repeatedly proven by DNA data from conidial and ascospore cultures.
Several Melanconium species have been described, which
could represent the conspicuous melanconium-like anamorph of M. spodiaea. However, as the original descriptions
are very brief and incomplete and as original material is
often unavailable for study, these cannot be confidently
connected to the currently accepted Melanconiella species.
In addition, they often consist of mixed elements considering
the various hosts listed, and most of these binomials have
not been used in recent literature. They should therefore be
considered dubious names.
Key to species of Melanconiella
Note: Although the anamorph is often important and helpful in the distinction of similar species and is therefore essential for
identification, this feature has to be evaluated carefully in species growing on Carpinus betulus. In the present study two
different species were often found on the same twig, e.g. species with hyaline and dark brown conidia: M. carpinicola/M.
chrysomelanconium and M. chrysomelanconium/M. chrysodiscosporina; or M. hyperopta/M. chrysodiscosporina, both with
hyaline conidia but with different conidial shapes and contents. Ascospores and conidia should be examined in water mounts,
as in KOH some features are either quickly disappearing (appendages, hyaline sheath) or less evident (swelling of ascospore
walls).
1 Ascospores dark brown; conidia dark brown .................................................................................................................................... 2
Ascospores hyaline; conidia hyaline or dark brown ............................................................................................................................ 3
2(1) Ascospores with hyaline, broad cap-like appendages; conidia usually not distinctly pip-shaped; on Carpinus in
Europe ......................................................................................................................................................... M. spodiaea
Ascospores without appendages; conidia pip-shaped; on Betula spp. in Europe and North America ....................... M. decorahenis
3(1) On Carpinus ................................................................................................................................................................................ 4
On other hosts ................................................................................................................................................................................... 12
4(3) Ascospores with distinctly swelling ascospore wall and with monomorphic, (sub)globose to bullet-shaped ascospore
cells mostly wider than 7 μm; commonly with bright yellow ectostroma; in Europe .................................................. 5
Ascospores without distinctly swelling ascospore wall and with slightly to distinctly dimorphic cells mostly narrower than 7 μm;
ectostroma grey, brownish, cinnamon, cream or yellow ............................................................................................................ 7
5(4) Ascospores (16–)19–23(–27.5)×(7.5–)9–12.5(–15.5) μm; always associated with conspicuous conidiomata containing dark
brown conidia; on C. betulus in open habitats (parks, hedges, forest edges) ........................................ M. chrysomelanconium
Ascospores (15.5–)17–20(–22)×(6–)7–10(–15) μm; with conidiomata containing hyaline conidia ................................................. 6
6(5) On C. betulus; conidia (12.5–)13.5–16(–19)×4.5–5.5(–6) μm, l/w0(2.2–)2.5–3.3(–4) ..........................M. chrysodiscosporina
On C. orientalis; conidia (11.5–)12.5–14(–15.5)×(5.5–)6–7(–7.5) μm, l/w0(1.7–)1.9–2.2(–2.7) ........................ M. chrysorientalis
7(4) Ascospores mostly wider than 5.5 and longer than 18.5 μm .................................................................................................... 8
Ascospores mostly narrower than 5.5 μm and shorter than 18.5 μm ............................................................................................. 10
Fungal Diversity (2012) 57:1–44
41
8(7) On Carpinus in Europe; ectostromatic discs concave, flat or slightly pustulate and little projecting above the perithecial level,
whitish to dull yellow or brownish; central column pale yellowish to pale greyish brown or brownish, entostroma inconspicuous, pale, slightly lighter than bark interior ............................................................................................................................... 9
On C. caroliniana in North America; ectostromatic discs well-developed, often pulvinate, commonly with distinctly
projecting ostioles thus appearing spiny, cream yellow, honey, cinnamon, orange brown to brown; central column
young bright olive-yellow, then olive-green to olive-brown; entostroma well-developed, of compacted hyphae, olivegreen or olive-brown ........................................................................................................................................ M. echinata
9(8) On C. betulus ........................................................................................................................................................... M. hyperopta
On C. orientalis ........................................................................................................................................ M. hyperopta var. orientalis
10(7) On C. betulus in Europe; in open habitats (parks, hedges, forest edges) .......................................................... M. carpinicola
On C. caroliniana in North America ................................................................................................................................................ 11
11(10) Ectostromatic disc mostly irregularly shaped, commonly surrounded by irregular edges or flaps of bark, drab, light to dark
grey or dark brown; ostioles mostly marginal in the disc; central column grey or brownish ....................................... M. ellisii
Ectostromatic discs regularly shaped, mostly circular, angular or ellipsoid, distinctly projecting, often pulvinate, pale or bright yellow
to light brown; ostioles mostly regularly arranged in the disc; central column yellow, olive grey or brownish ............... M. elegans
12(3) On Corylus; ascospores broadly fusoid; ascospore ends distinctly subacute with persistent knob-like hyaline
appendages .................................................................................................................................................... M. flavovirens
On Ostrya; ascospores fusoid; ascospore ends narrowly rounded to subacute, without or with cap-like appendages ................... 13
13(12) Ascospores (20–)23–28(–37.5)×(5–)6.5–8(–9.5) μm; conidia hyaline; on Ostrya carpinifolia in Europe ......... M. meridionalis
Ascospores (14.5–)16.5–19(–23)×(4–)5.5–7(–8.5) μm; with dark brown conidia having a light brown equatorial zone; on Ostrya
virginiana in North America ......................................................................................................................................... M. ostryae
Species excluded from Melanconiella
Apart from the species of Melanconiella recognised in the
current study, most names described or combined in the
genus are either incompletely known or doubtful members
of Melanconiella. As we have not been able to study material of some taxa, the list below contains only species for
which we have some evidence. The account also includes
taxa that were previously synonymised with M. chrysostroma (fide Wehmeyer 1941) and now found to be unrelated
to Melanconiella after studying their types.
Diatrype carpinigera Berk. & M.A. Curtis, Grevillea 4:96
(1876).
Synonym: Melanconis carpinigera (Berk. & M.A. Curtis)
Petr., Sydowia 6:355 (1952).
Based on a rudimentary description, this name was combined into Diaporthe by Saccardo (1883), subsequently
synonymised with Melanconis chrysostroma var. ellisii by
Wehmeyer (1941) and combined into Melanconis by Petrak
(1952b). Re-examination of the type (K(M) 171592)
showed that it is not a diaporthalean fungus; stroma morphology and shape and size (18.5–22.5×8.5–11.5 μm) of its
brown, two-celled ascospores rather suggest a Valsaria sp.
Melanconiella appendiculata (G.H. Otth) Sacc., Syll Fung
11:XXIX (1895).
This species is currently classified as Prosthecium appendiculatum (G.H. Otth) M.E. Barr and is phylogenetically
unrelated to Melanconiella (unpubl. data).
Melanconiella chrysostroma (Fr.) Sacc., Syll Fung 1:741
(1882).
Basionym: Valsa chrysostroma Fr., Sum Veg Scand:412
(1849).
[0 Sphaeria xanthostroma Mont. & Fr. (1834), non
Schwein. (1832), non J.C. Schmidt (1817)].
Synonym: Melanconis chrysostroma (Fr.) Tul., Selecta
fung carpol (Paris) 2:125 (1863).
Melanconiella chrysostroma is based on Valsa chrysostroma, which was established by Fries (1849) as a new
name for the homonym Sphaeria xanthostroma Mont. & Fr.
(Montagne 1834), non Schwein. (von Schweinitz 1832).
However, Sphaeria xanthostroma Mont. & Fr. is the basionym of Wuestneia xanthostroma (Reid and Booth 1989) and
represents a taxon differing from Melanconiella by unicellular ascospores. No morphological description was provided by Fries (1849). Tulasne and Tulasne (1863), who
apparently did not know Wuestneia xanthostroma, considered the description and illustrations of Montagne (1834: pl.
12, Fig. 4) to be unreliable (“de peritheciorum forma et
thecarum fecunditate non fida”), misapplied the name for a
taxon with two-celled ascospores and combined the name
into Melanconis, explicitly citing Sphaeria xanthostroma
Mont. as a synonym and therefore following the nomenclatural concept of Fries (1849). Because the basionym, Valsa
chrysostroma, is based on the type and description of
Sphaeria xanthostroma Mont., Melanconis chrysostroma is
an obligate synonym of Wuestneia xanthostroma, although
the concept of Tulasne and Tulasne (1863) applies to a
42
different taxon belonging to Melanconiella. Their description does not fit a single Melanconiella species, but is rather
based on a mixture of several taxa. Tulasne and Tulasne
(1863) record both brown, ovate conidia, 13–15×10 μm, as
well as biguttulate hyaline conidia, 10–13×3.5–5 μm. Their
descriptions, measurements and illustrations could be based
on a mixture of M. chrysodiscosporina (with hyaline biguttulate conidia) and M. chrysomelanconium (with brown
conidia), which agrees with the illustration of biguttulate
ascospores. However, they do not give ascospore sizes, but
state that they are of similar width as the hyaline conidia, and
they record a biseriate ascospore arrangement, features which
are not in line with the former two species, but rather imply
M. hyperopta or M. carpinicola. As we did not have the
opportunity to examine the Tulasne collections, their concept
of Melanconis chrysostroma remains obscure.
Melanconiella corni (Wehm.) Petr., Sydowia 6:15 (1952).
This species is currently classified as Hapalocystis corni
(Wehm.) M.E. Barr and is phylogenetically unrelated to
Melanconiella (unpubl. data).
Melanconiella leucostroma (Niessl & Rehm) Sacc., Syll
Fung 2: Addenda Vol I:LIV (1883).
According to Wehmeyer (1941) this name is a synonym
of Cryptodiaporthe galericulata.
Melanconiella meschuttii (Ellis & Everh.) Berl. & Voglino,
in Saccardo, Syll Fung, Addit. I–IV:129 (1886).
According to Wehmeyer (1941) this name is a synonym
of Melanconis nigrospora, now classified as Chapeckia
nigrospora (Peck) M.E. Barr.
Melanconiella nigrospora (Peck) Dearn. & House, Bull N
Y State Mus 266:81 (1925).
This is a synonym of Chapeckia nigrospora, which is
unrelated to Melanconiella (Fig. 1).
Melanconiella nyssaegena Ellis & Everh., J Mycol 8:18
(1902).
According to Wehmeyer (1941) this name is a synonym
of Massariovalsa sudans (Berk. & M.A. Curtis) Sacc.
Melanconiella obruta (Ellis & Everh.) Sacc. & P. Syd., Syll
Fung 16:509 (1902).
According to Wehmeyer (1941) this name is a nomen
dubium which cannot be interpreted because the type collection contains only a melanconium-like anamorph.
Melanconis platystroma Wehm., Mycologia 29:613 (1937).
Melanconis platystroma was described from Carpinus
caroliniana in Michigan and included in Wehmeyer
Fungal Diversity (2012) 57:1–44
(1941). Although it may belong in Melanconiella, it is
unlike any species included here and differs in having truncated, grey to yellow grey ectostromatic discs with scarcely
visible or barely erumpent ostioles, large, brown ascospores
(25–31×12.5–14.5 μm) with a distinctly greenish tint when
fresh, and large (20–31 × 16–18 μm), brown conidia
(Wehmeyer 1937, 1941). Wehmeyer (1937) noted that he
was unable to germinate the ascospores and conidia despite
numerous attempts, which is atypical for species of
Melanconiella, conidia and ascospores of which germinate
easily in pure culture. Wehmeyer (1941) noted that its stromata closely match those of Massariovalsa sudans which,
according to our experience, is also difficult to grow.
Apparently known only from the type collection, this species cannot be appropriately classified until fresh material
becomes available.
Melanconiella stellata Petch, Ann Roy Bot Gard Peradeniya
7:306 (1922).
According to Wehmeyer (1941) this name may be a
species of Valsaria.
Valsa bitorulosa Berk. & Br., Ann Mag Nat Hist Ser 3,
3:367 (1846).
Synonym: Diaporthe bitorulosa (Berk. & Broome) Sacc.,
Syll Fung 1: 608 (1882).
Based on a scanty description, this name was combined
into Diaporthe by Saccardo (1882) and subsequently synonymised with Melanconis chrysostroma by Wehmeyer (1937,
1941). Re-examination of the type (K(M) 171593) showed
that it is not a diaporthalean fungus; stroma morphology
and shape and size (19–20×5.5–7 μm) of its brown,
four-celled ascospores rather suggest a Thyridaria sp.
Acknowledgements We thank Walter Gams for hospitality and excursion support in Italy; Jacques Fournier, Enrique Rubio Domínguez,
Sven-Åke Hanson and Larissa Vasilyeva for collecting and communicating Melanconiella specimens; Irmgard Greilhuber and her family
for organising and participating in numerous collecting trips together
with HV; the fungarium curators of B, BPI, DAOM, G, GZU, K, M,
NY, UPS and W for the loan of specimens; Scott Redhead (DAOM) for
providing notes of L. E. Wehmeyer and for allowing DNA extraction
from the type specimen of M. echinata; Walter Till (WU) for managing
the herbarium loans; and the British Mycological Society for invitation
to the BMS Spring Foray 2011 in Yorkshire.
References
Barr ME (1978) The Diaporthales in North America, with emphasis on
Gnomonia and its segregates. Mycol Mem 7:1–232
Castlebury LA, Rossman AY, Jaklitsch WJ, Vasilyeva LN (2002) A
preliminary overview of the Diaporthales based on large subunit
nuclear ribosomal DNA sequences. Mycologia 94:1017–1031
Fungal Diversity (2012) 57:1–44
Chaverri P, Samuels GJ (2003) Hypocrea/Trichoderma (Ascomycota,
Hypocreales, Hypocreaceae): species with green ascospores. Stud
Mycol 48:1–116
Clements FE, Shear CL (1931) Genera of fungi, 2nd edn. H.W. Wilson,
New York
de Hoog GS, Gerrits van den Ende AHG (1998) Molecular diagnostics
of clinical strains of filamentous basidiomycetes. Mycoses
41:183–189
De Silva H, Castlebury LA, Green S, Stone JK (2009) Characterisation
and phylogenetic relationships of Anisogramma virgultorum and
A. anomala in the Diaporthales (Ascomycota). Mycol Res
113:73–81
Dennis RWG (1968) British Ascomycetes. J. Cramer, Stuttgart
Druzhinina IS, Kubicek CP, Komoń-Zelazowska M, Mulaw TB, Bissett J
(2010) The Trichoderma harzianum demon: complex speciation
history resulting in coexistence of hypothetical biological species,
recent agamospecies and numerous relict lineages. BMC Evol Biol
10:94
Edgar RC (2004) MUSCLE: multiple sequence alignment with high
accuracy and high throughput. Nucleic Acids Res 32:1792–1797
Ellis MB, Ellis JP (1997) Microfungi on land plants. An identification
handbook. Richmond Publishing, Slough
Ellis JB, Everhart BM (1892) North American pyrenomycetes. Ellis &
Everhart, Newfield
Fries EM (1849) Summa Vegetabilium Scandinaviae. Sectio posterior.
259–572. A. Bonnier, Stockholm, Leipzig
Fuckel KWGL (1870) Symbolae Mycologicae. Beiträge zur Kenntnis
der Rheinischen Pilze. Jahrb Nassau Ver Naturkd 23–24:1–459
Fuckel KWGL (1871) Symbolae Mycologicae. Erster Nachtrag. Jahrb
Nassau Ver Naturkd 25–26:289–346
Fuckel KWGL (1874) Symbolae Mycologicae. Zweiter Nachtrag.
Jahrb Nassau Ver Naturkd 27–28:1–99
Gazis R, Rehner S, Chaverri P (2011) Species delimitation in fungal
endophyte diversity studies and its implications in ecological and
biogeographic inferences. Mol Ecol 20:3001–3013
Gerhardt E, Hein B (1979) Die nomenklatorischen Typen der von Th.
Nitschke beschriebenen Arten im Pilzherbar des Botanischen
Museums Berlin-Dahlem. Willdenowia 9:313–329
Hall TA (1999) BioEdit: a user-friendly biological sequence alignment
editor and analysis. program for Windows 95/98/NT. Nucleic
Acids Symp Ser 41:95–98
Huelsenbeck JP, Ronquist F (2001) MRBAYES: Bayesian inference of
phylogenetic trees. Bioinformatics 17:754–755
Jaklitsch WM (2009) European species of Hypocrea part I. The greenspored species. Stud Mycol 63:1–91
Jaklitsch WM (2011) European species of Hypocrea part II. species
with hyaline ascospores. Fungal Divers 48:1–250
Jaklitsch WM, Voglmayr H (2004) Hapalocystis occidentalis - a new
species of Diaporthales from North America and a key to the
species of Hapalocystis. Stud Mycol 50:229–234
Jaklitsch WM, Voglmayr H (2011) Nectria eustromatica sp. nov., an
exceptional species with a hypocreaceous stroma. Mycologia
103:209–218
Jaklitsch WM, Komon M, Kubicek CP, Druzhinina IS (2006) Hypocrea voglmayrii sp. nov. from the Austrian Alps represents a new
phylogenetic clade in Hypocrea/Trichoderma. Mycologia
97:1365–1378 [‘2005’]
Jaklitsch WM, Stadler M, Voglmayr H (2012) Blue pigment in Hypocrea caerulescens sp. nov. and two additional new species in sect.
Trichoderma. Mycologia 104(4):17
Katoh K, Toh H (2008) Recent developments in the MAFFT
multiple sequence alignment program. Brief Bioinform 9:286–
298
Katoh K, Misawa K, Kuma K, Miyata T (2002) MAFFT: a novel
method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Res 30:3059–3066
43
Kobayashi T (1970) Taxonomic studies of Japanese Diaporthaceae
with special reference to their life-histories. Bull Gov Forest Exp
Sta 226:1–242
Liu YL, Whelen S, Hall BD (1999) Phylogenetic relationships among
ascomycetes: evidence from an RNA polymerase II subunit. Mol
Biol Evol 16:1799–1808
Mejía LC, Castlebury LA, Rossman AY, Sogonov MV, White JF
Jr (2011a) A systematic account of the genus Plagiostoma
(Gnomoniaceae, Diaporthales) based on morphology, hostassociations, and a four-gene phylogeny. Stud Mycol 68:211–
235
Mejía LC, Rossman AY, Castlebury LA, White JF Jr (2011b) New
species, phylogeny, host-associations and geographic distribution
of genus Cryptosporella (Gnomoniaceae, Diaporthales). Mycologia 103:379–399
Montagne C (1834) Notice sur les plantes cryptogames récemment
découvertes en France, contenant aussi l’indication précise des
localités de quelques espèces les plus rares de la flore Française.
Ann Sci Nat Bot Sér 2(1):295–307
Müller E, von Arx JA (1962) Die Gattungen der didymosporen Pyrenomyceten. Beitr Kryptogamenfl Schweiz 11(2):1–922
Munk A (1957) Danish pyrenomycetes. A preliminary flora. Dansk
Bot Ark 17:1–491
Nylander JA, Wilgenbusch JC, Warren DL, Swofford DL (2008)
AWTY (are we there yet?): a system for graphical exploration
of MCMC convergence in Bayesian phylogenetics. Bioinformatics 24:581–583
Otth G (1869) Sechster Nachtrag zu dem in Nr. 15–23 der Mittheilungen
enthaltenen Verzeichnisse schweizerischer Pilze. Mitth Natforsch
Ges Bern 1868:37–70
Pavlic D, Slippers B, Coutinho TA, Wingfield MJ (2009) Multiple
gene genealogies and phenotypic data reveal cryptic species of the
Botryosphaeriaceae: a case study on the Neofusicoccum parvum ⁄
N. ribis complex. Mol Phyl Evol 51:259–268
Petrak F (1952a) Fungi beltsvillenses. III. Sydowia 6:5–16
Petrak F (1952b) Fungi beltsvillenses. IV. Sydowia 6:352–360
Petrak F (1962) Mykologische Beiträge zur österreichischen Flora.
Sydowia 16:155–198
Reid J, Booth C (1989) On Cryptosporella and Wuestneia. Can J Bot
67:879–908
Riethmüller A, Voglmayr H, Göker M, Weiß M, Oberwinkler F (2002)
Phylogenetic relationships of the downy mildews (Peronosporales) and related groups based on nuclear large subunit ribosomal
DNA sequences. Mycologia 94:834–849
Saccardo PA (1876) Fungi Veneti novi vel critici. Series V. Nuovo
Giornale Botanico Italiano 8:161–211
Saccardo PA (1882) Sylloge Fungorum 1. Padova: published by the
author
Saccardo PA (1883) Sylloge Fungorum 2. Padova: published by the
author
Saccardo PA (1884) Sylloge Fungorum 3. Padova: published by the
author
Schoch CL, Seifert KA, Huhndorf S, Robert V, Spouge JL, Levesque
CA, Chen W et al. (2012) Nuclear ribosomal internal transcribed
spacer (ITS) region as a universal DNA barcode marker for
Fungi. Proc Natl Acad Sci USA, www.pnas.org/cgi/doi/10.1073/
pnas.1117018109
Sieber TN (2007) Endophytic fungi in forest trees: Are they mutualists? Fungal Biol Rev 21:75–89
Silvestro D, Michalak I (2011) raxmlGUI: a graphical front-end for
RAxML. Org Divers Evol. doi:10.1007/s13127-011-0056-0
Stamatakis E (2006) RAxML-VI-HPC: maximum likelihood-based
phylogenetic analyses with thousands of taxa and mixed models.
Bioinformatics 22:2688–2690
Sutton BC (1980) The Coelomycetes. Commonwealth Mycological
Institute, Kew
44
Swofford DL (2002) PAUP*: Phylogenetic Analysis Using Parsimony
(*and Other Methods), Version 4.0b10. Sinauer Associates, Sunderland, Massachusetts
Thiers B (2012) Index Herbariorum: a global directory of public
herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/ih/
Tulasne LR, Tulasne C (1863) Selecta Fungorum Carpologia, vol. 2.
Paris
Vasilyeva LN, Stephenson SL (2010) Biogeographical patterns in
pyrenomycetous fungi and their taxonomy. 1. The Grayan disjunction. Mycotaxon 114:281–303
Vilgalys R, Hester M (1990) Rapid genetic identification and mapping
of enzymatically amplified ribosomal DNA from several Cryptococcus species. J Bacteriol 172:4238–4246
Voglmayr H, Jaklitsch WM (2008) Prosthecium species with Stegonsporium anamorphs on Acer. Mycol Res 112:885–905
Voglmayr H, Jaklitsch WM (2011) Molecular data reveal high host
specificity in the phylogenetically isolated genus Massaria (Ascomycota, Massariaceae). Fungal Divers 46:133–170
Fungal Diversity (2012) 57:1–44
von Höhnel F (1918) Mycologische Fragmente. Ann Mycol 16:35–174
von Schweinitz LD (1832) Synopsis fungorum in America boreali
media degentium. Trans Am Philos Soc II 4:141–316
Wehmeyer LE (1926) Cultural life histories of Melanconis and Pseudovalsa. Mycologia 18:257–273
Wehmeyer LE (1933) The genus Diaporthe Nitschke and its segregates. Univ Michigan Stud Sci Ser 9:1–349
Wehmeyer LE (1937) Studies of certain species of Melanconis on
Carpinus, Ostrya and Corylus. Mycologia 29:599–617
Wehmeyer LE (1941) A revision of Melanconis, Pseudovalsa, Prosthecium and Titania. Univ Michigan Stud, Sci Ser 14:1–161
Werle E, Schneider C, Renner M, Völker M, Fiehn W (1994) Convenient single-step, one tube purification of PCR products for direct
sequencing. Nucleic Acids Res 22:4354–4355
White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct
sequencing of fungal ribosomal RNA genes for phylogenetics. In:
Innis MA, Gelfand DH, Sninsky JJ, White TJ (eds) PCR protocols: a guide to methods and applications. Academic, San Diego,
pp 315–322