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Phylogenetic insights resolve Dacampiaceae (Pleosporales)
as polyphyletic: Didymocyrtis (Pleosporales, Phaeosphaeriaceae)
with Phoma-like anamorphs resurrected and segregated
from Polycoccum (Trypetheliales,Polycoccaceae fam. nov.)
Damien Ertz
1
&Paul Diederich
2
&James D. Lawrey
3
&Franz Berger
4
&
Colin E. Freebury
5
&Brian Coppins
6
&Alain Gardiennet
7
&Josef Hafellner
8
Received: 2 June 2015 /Accepted: 20 August 2015
#School of Science 2015
Abstract A phylogenetic analysis of nuLSU and ITS se-
quences representing genera previously included in
Dacampiaceae indicates that the family is strongly poly-
phyletic and that the type species of Dacampia is placed
in Pleosporales. The genus Munkovalsaria s. str. is placed
in Didymosphaeriaceae (Pleosporales). Polycoccum s. str.
and two species of Clypeococcum are shown to form a
new lineage sister to the Trypetheliaceae in Trypetheliales
and described here as Polycoccaceae. Other members of
Polycoccum s. lat. are included in the Pleosporales and
are closely related to lichenicolous Phoma-like species of
the family Phaeosphaeriaceae. The genus Didymocyrtis is
resurrected for these species and for lichenicolous species
previously assigned to Diederichia,Diederichomyces,
Leptosphaeria and Phoma. The genera Diederichia and
Diederichomyces are synonymized with Didymocyrtis.
The new c o m b i n a t i ons D i d y m o c y r t is bryonthae,
D. cladoniicola,D. foliaceiphila,D. infestans,
D. kaernefeltii,D. mela ne lixiae,D. pseudeverniae,
D. ramalinae,D. slaptoniensis and D. xanthomendozae
are made, and the new name D. epiphyscia is introduced
for Phoma physciicola. Some anamorph-teleomorph rela-
tionships are resolved, such as Didymocyrtis ramalinae–
Phoma ficuzzae and Didymocyrtis consimilis–Phoma
caloplacae, the phylogenetic results being supported by
single ascospore cultures that lead to the asexual stage
producing pycnidia and conidia in culture. Speciation by
host switching is assumed to be important in the genus
Didymocyrtis. An identification key to Didymocyrtis spe-
cies is provided.
*Damien Ertz
damien.ertz@br.fgov.be
Paul Diederich
paul.diederich@education.lu
James D. Lawrey
jlawrey@gmu.edu
Franz Berger
flechten.berger@gmail.com; flechten.berger@aon.at
Colin E. Freebury
cfreebury@mus-nature.ca
Brian Coppins
lichensel@btinternet.com
Alain Gardiennet
agardiennet@gmail.com
Josef Hafellner
josef.hafellner@uni-graz.at
1
Department Bryophytes-Thallophytes (BT), Botanic Garden Meise,
Nieuwelaan 38, B-1860 Meise, Belgium
2
Musée national d’histoire naturelle, 25 rue Munster,
L-2160 Luxembourg, Luxembourg
3
Department of Biology, George Mason University, 4400 University
Drive, Fairfax, VA 22030, USA
4
Raiffeisenweg 130, A-4794 Kopfing, Austria
5
Canadian Museum of Nature, PO Box 3443, Stn. BD^,
Ottawa, Ontarion K1P 6P4, Canada
6
Royal Botanic Garden Edinburgh, 20A Inverleith Row,
Edinburgh EH3 5LR, UK
7
14 rue Roulette, F-21260 Véronnes, France
8
Institute for Plant Sciences, NAWI Graz, Karl-Franzens-University
Graz, Holteigasse 6, A-8010 Graz, Austria
Fungal Diversity
DOI 10.1007/s13225-015-0345-6
Keywords Dothideomycetes .Leptosphaeria .Diederichia .
Diederichomyces .Lichenicolous fungi .Phylogeny
Introduction
The family Dacampiaceae Körb. includes lichenicolous,
lichenized and saprobic fungi forming blackish, perithecioid
ascomata, a pseudoparenchymatous exciple, branched-
anastomosing pseudoparaphyses, bitunicate asci, and septate
to muriform, usually brown ascospores (Hyde et al. 2013).
The family was placed in Dothideales in the past (Eriksson
and Hawksworth 1986,1993; Hawksworth and Diederich
1988; Henssen 1995), but is currently considered as
‘Dothideomycetes, family incertae sedis’(Hyde et al. 2013;
Wijayawardene et al. 2014). Dacampiaceae is considered to
be a heterogeneous assemblage including about 110 species in
10 genera: A aosphae ria ,Cly peococc um,D aca mpi a,
Eopyrenula,Leptocucurthis,Munkovalsaria,Polycoccum,
Pseudonitschkia,Pyrenidium and Weddellomyces (Hyde
et al. 2013).
Few sequences of Dacam pia cea e are available in
GenBank. One sequenced species is M unk oval sar ia
appendiculata Aptroot, collected on dead culms of Zea mays
and assigned to Pleosporales in a phylogenetic analysis by
Apt r o ot (2004); it als o w a s f ound to cluster wit h
Montagnula opulenta (Didymosphaeriaceae as recently
circumscribed by Ariyawansa et al. 2014) in a phylogeny by
Wang et al. (2007). Sequences of M. donacina (generic type)
were generated by Pitt et al. (2014), but not included in a
phylogenetic an a l y s i s . A 28 S r D N A s e quence of
Polycoccum vermicularium was also deposited in GenBank
(AY961601), but we found it to be of a poor quality and its
reliability is questioned here.
Many de s c ribe d s p ecies o f D a ca m p i a cea e ar e
lichenicolous, but limited sequence data for lichenicolous spe-
cies means they are not commonly included in cladograms
based on molecular data sets (e.g., Lutzoni et al. 2004;
Zhang et al. 2009). Despite some recent progress in placing
lichenicolous species in Dothideomycetes (Lawrey et al. 2011,
2012; Ruibal et al. 2011; Ertz et al. 2013; Pérez-Ortega et al.
2013; Ertz and Diederich 2015), questions remains about
whether the placement and content of Dacampiaceae, includ-
ing lichenicolous species, are correct.
Polycoccum, the most species-rich genus of Dacampiaceae
and one that includes only lichenicolous species, has never
been adequately studied using molecular data and is here stud-
ied in detail. Polycoccum had been little used for lichenicolous
ascomycetes until its reinstallation by Santesson (1960), but in
the past two decades it has become one of the most species-
rich genera of lichenicolous fungi, with 57 species accepted
by Lawrey and Diederich (2015) and new species described
almost every year. A first key was given by Vězda (1969, 10
species). The most complete key so far available is that com-
piled by Hawksworth and Diederich (1988, 23 species world-
wide), and there are additional more recently-published re-
gional keys (e.g., Atienza et al. 2003, 13 species from Spain;
Calatayud 2004, 5 species from the Sonoran desert region;
Ihlen and Wedin 2008, 14 species from Sweden; Gardiennet
2012, 13 species from France) that illustrate the diversity of
this genus in various parts of the world.
The relationship of Polycoccum to other members of the
Dothideales has never been established with certainty. Crivelli
(1983: 193) suggested a close relationship among Dacampia,
Polycoccum and Pyrenidium (sub Dacampiosphaeria). This
view was further elaborated by Hawksworth and Diederich
(1988), when they proposed to treat these three genera togeth-
er with Byssothecium,Clypeococcum and Weddellomyces in
the family Dacampiaceae in the Dothideales. However, until
now these hypotheses have not been adequately tested.
This paper aims at (1) testing the monophyly of the family
Dacampiaceae, (2) elucidating the phylogenetic position of
the genera Clypeococcum,Diederichia and Polycoccum, (3)
testing the monophyly of the genus Polycoccum, (4) reinstat-
ing the genus Didymocyrtis,and (5) testing some anamorph-
teleomorph relationships among lichenicolous members of the
genera Polycoccum s. lat. and Phoma s. lat.
Material and methods
Morphological study
Herbarium specimens are deposited in BR, CANL, E, GZU,
M, NY, UBC and in the private collections of S. Beeching, F.
Berger, P. Diederich, J. Hafellner and J. Etayo. External mor-
phology of herbarium specimens was examined and measured
under Leica MZ 7.5 (magnification up to ×50), Wild M3
(magnification up to ×40) and Olympus SZX12 (magnifica-
tion up to ×90) stereomicroscopes. Macroscopic photographs
were done using a Canon 40D camera with a Canon MP-E
65 mm lens or a Nikon BD Plan 10× microscope objective,
StackShot (Cognisys) and Helicon Focus (HeliconSoft) for
increasing the depth of field; or with a Color View I digital
camera connected to an Olympus SZX12 stereomicroscope.
Hand-made sections of ascomata and thallus were studied in
water, 5 % KOH (K), or 1 % I
2
Lugol’s reagent without (I) or
with KOH pre-treatment (K/I) using Leica DMLB, Leica
DMRE and Olympus BX51 microscopes (magnification up
to ×1000). Sectioning was performed by one of us (JH) using
a freezing microtome (Leitz, sections of 12–15 μm), but
squash preparations were also used, especially for ascus anal-
ysis. Conidiogenesis of conidia was studied in water and
erythrosin B (ALDRICH 19,826-9) in 10 % ammonia.
Measurements of asci, ascospores and conidia all refer to ma-
terial examined in tap water. Conidial size of a large number of
Fungal Diversity
conidia was measured for some specimens, and the average
(X) and standard deviation (SD) calculated. Such measure-
ments are given as X ± SD, surrounded by the extreme values
(between parentheses), followed by the number of measure-
ments (n). For the graphical representation of the conidial size
distribution (Fig. 6), each specimen is represented by an ellip-
se, of which the centre represents the average length and
breadth, the large radius represents the SD of conidial length
and the small radius the SD of conidial breadth. The graphical
representation has been done using the PostScript language.
Microscopic photographs were prepared using either a Leica
DMLB microscope with differential interference contrast and
fitted with a Leica EC3 camera, or an Olympus BX51 micro-
scope with interference contrast and connected to a Color
View I digital camera.
Molecular techniques
Cultures were isolated from ascospores (single ascospore cul-
tures for sexual stages of Didymocyrtis ramalinae, for
Didymocyrtis aff. consimilis Ertz 17617b, Berger 26876, and
for Didymocyrtis slaptoniensis MoraA-B; two-ascospores iso-
late for Didymocyrtis consimilis 12041; and multi-ascospores
isolate for Didymocyrtis melanelixiae and D. slaptoniensis
12009), or from conidia (multiconidia cultures) offreshly col-
lected material on malt-yeast extract medium as described by
Yoshimura et al. (2002) (Fig. 9). Thin sections were made
through ascomata or pycnidia, and the outer wall was removed
with a sterile razor blade to expose ascospores or conidia,
which were then spread directly on malt-yeast extract agar.
Isolated ascospores were immediately transferred to new petri
dishes to attain single-spore cultures. Germination of conidia
and ascospores was often observed within a day. The cultures
were kept at room temperature in the laboratory of the Botanic
Garden Meise and exposed to a natural day-light regime.
Cultures maintained at George Mason University were kept
at 18 °C in 12 h light-12 h dark cycles. No experiments were
done to test whether different light or temperature conditions
could improve the growth rate. All the strains were fast-
growing and therefore only a few weeks were required to
obtain sufficient material for DNA extraction. In some cases,
hand-made sections of the hymenium or thallus were used for
direct PCR as described in Ertz et al. (2014). The outer wall of
ascomata was removed with a sterile razor blade to isolate the
hymenium. The material was then added to a tube containing
the PCR reaction mixture and amplified directly. Genomic
DNA was isolated from cultures using the CTAB extraction
protocol (Doyle and Doyle 1990). Amplification reactions
were prepared for a 50 μl final volume containing 5 μl 10×
DreamTaq Buffer (Fermentas), 1.25 μl of each of the 20 μM
primers, 5 μl of 2.5 mg mL
−1
bovin serum albumin
(Fermentas #B14), 4 μl of 2.5 mM each dNTPs (Fermentas),
1.25 U DreamTaq DNA polymerase (Fermentas), and 1 μl of
template genomic DNA or tiny fragments of fungal material.
A targeted fragment of about 1.1 kb at the 5′end of the nuLSU
rDNA was amplified using primers LIC15R (Miadlikowska
et al. 2002) with LR6 (Vilgalys and Hester 1990). A fragment
of about 0.6 kb of the nuITS rDNA was amplified using
primers ITS1F and ITS4 (White et al. 1990). The yield of
the PCRs was verified by running the products on a 1 %
agarose gel using ethidium bromide. Both strands were se-
quenced by Macrogen® using amplification primers.
Additional primers were used for the sequencing of nuLSU:
LR3, more rarely LR3R and LR5 (Vilgalys and Hester 1990)
(Vilgalys’website, http://www.botany.duke.edu/fungi/
mycolab). Sequence fragments were assembled with
Sequencher version 4.6 (Gene Codes Corporation, Ann
Arb o r , M ichi g an) . S eque n ces w e re su b ject e d to
MEGABLAST searches to verify their closest relatives and
to detect potential contaminations.
Taxon selection and phylogenetic analyses
Although not possible for all taxa, we tried to achieve a sample
series with at least two specimens of each newly sequenced
species, preferably from different localities, to avoid misiden-
tifications caused by contamination or sequencing errors. Our
sample series include sequences obtained from the type spe-
cies of Dacampia,Didymocyrtis and Polycoccum that are im-
portant for taxonomic conclusions. For the nuLSU phyloge-
netic tree, the closest relatives of the new sequences based on
BLAST searches were retrieved from GenBank. Additional
taxa were selected from Hyde et al. (2013) and Nelsen et al.
(2014) to include a wide array of families belonging to
Dothideomycetes and an exhaustive list of genera belonging
to Trypetheliales. The original nuLSU matrix used in Ertz and
Diederich (2015) was used as main template. The resulting
nuLSU matrix consisted of 109 sequences, mainly from a
wide variety of Dothideomycetes. Some Arthoniomycetes
were also included. Three outgroup species were chosen to
represent the class Eurotiomycetes (Caliciopsis pinea and
Capronia munkii) and the class Leotiomycetes (Lachnum
virgineum). Caliciopsis pinea was used as the rooting taxon
in the related analyses. The ITS matrix was assembled manu-
ally and consisted of 58 sequences, including 52 sequences of
Didymocyrtis and 6 sequences chosen from the most closely
related gener a in Tr akunyingcharoen et al. (2 014).
Phaeosphaeria lycopodina was used as the rooting taxon in
these analyses. The alignments were improved manually
using MacClade 4.05 (Maddison and Maddison 2002).
Terminal ends of sequences, ambiguous aligned regions and
introns were delimited manually and excluded from the
nuLSU dataset. The nuLSU data set consisted of 1160 unam-
biguously aligned characters, of which 463 were variable. The
Fungal Diversity
ITS data set consisted of 594 unambiguously aligned charac-
ters, of which 120 were variable.
The best-fit model of DNA evolution GTR+I+G was cho-
sen for the nuLSU data set and the GTR+I for the ITS data set
using the Akaike information criterion (AIC; Akaike 1973) as
implemented in Modeltest v. 3.7 (Posada and Crandall 1998).
Bayesian analyses were carried out using the Metropolis-
coupled Markov chain Monte Carlo method (MCMCMC) in
MrBayes v. 3.2.3 (Huelsenbeck and Ronquist 2001; Ronquist
and Huelsenbeck 2003) on the CIPRES portal (Miller et al.
2010). Analyses were run under the selected model of nucle-
otide substitution with six rate categories. Two parallel
MCMCMC runs were performed, with each run using four
independent chains and 100,000,000 generations for the
nuLSU data set and for the ITS data set, and sampling trees
every 1000th generation. Convergence diagnostics were esti-
mated using the PSRF (Potential scale reduction factor) where
values closer to one indicated convergence between runs
(Gelman and Rubin 1992), and using TRACER v. 1.6 by
plotting the log-likelihood values of the sample points against
generation time (Rambaut and Drummond 2007). Posterior
probabilities (PP) of the nuLSU and ITS matrices were deter-
mined by calculating a majority-rule consensus tree generated
from the 150,002 post-burnin trees of the 200,002 trees sam-
pled by the two MCMCMC runs using the sumt option of
MrBayes. In addition, a Maximum Likelihood (ML) analysis
was performed on the nuLSU and ITS data sets using GARLI
(Zwickl 2006, v. 0.951 for OSX) with default settings, and a
single most likely tree was produced (–lnL=11690.8948 for
the nuLSU tree; –lnL=1941.3522 for the ITS tree). One thou-
sand bootstrap pseudoreplicates were used to calculate a ma-
jority rule consensus tree in PAUP* 4.0b10 (Swofford 2002)
to assess the Maximum Likelihood bootstrap values (ML-bs).
ML-bs≥70 % and PP≥95 % were considered to be signifi-
cant. Phylogenetic trees were visualized using FigTree v. 1.4.2
(Rambaut 2012).
Results
We obtained 54 new sequences (20 nuLSU and 34 ITS) be-
longing to 15 species from Austria, Belgium, Canada, Canary
Islands (Gomera), Madeira, France (including Corsica),
Iceland, Luxembourg, Scotland, Switzerland and USA
(Table 1). The Bayesian nuLSU tree we recovered did not
contradict the ML nuLSU tree topology for the strongly sup-
ported branches and hence only the ML tree is shown with the
branches having ML-bs≥70 % in bold and with the posterior
probabilities of the Bayesian analysis added above the internal
branches (Fig. 1). The recovered Bayesian ITS tree did not
contradict the ML ITS tree’s topology for the strongly sup-
ported branches and hence only the ML tree is shown with the
branches having ML-bs≥70 % in bold and with the posterior
probabilities of the Bayesian analysis added above the internal
branches (Fig. 2).
The genera Clypeococcum, Dacampia, Diederichia (here
synonymized with Didymocyrtis, see Taxonomy section) and
Polycoccum are included in a molecular phylogeny for the first
time, all except the first being represented by the type species.
The status of Clypeococcum remains unclear until the type,
C. cladonema is sequenced. The backbone of our nuLSU phy-
logenetic tree is poorly resolved. However, main groups such as
Arthoniales,Capnodiales,Eremithallales,Hysteriales,
Trypetheliales,Tubeufiales and Venturiales are strongly support-
ed by both analyses, even though the relationships among such
groups are not supported. Polycoccum is resolved as polyphylet-
ic, with species being placed in two main distantly-related line-
ages corresponding to Trypetheliales (as newly defined) and
Pleosporales. Trypetheliales is strongly supported and divided
into two strongly supported main lineages corresponding to
Polycoccaceae (newly described here), and to Trypetheliaceae.
Polycoccaceae includes two genera, Clypeococcum and
Polycoccum, of which the latter is paraphyletic. Pleosporales is
only supported by the Bayesian analysis (PP=96).
Dacampia, the type genus of Dacampiaceae, is strongly
supported by both analyses (ML-bs=92, PP=100) and placed
in Pleosporales. It is sister to Paraleptosphaeria orobanches,
but this relationship is not supported. The relationship of
Dacampiaceae with other families of Pleosporales cannot
be resolved with certainty using only nuLSU sequences and
the available representatives of Pleosporales. Didymocyrtis is
nested in Pleosporales and includes eleven specimens that
form a poorly supported clade with very low resolution.
Figur e 2p r esen t s a m ore de t a iled ph y l ogeny o f
Didymocyrtis based on ML analysis of ITS sequence data.
According to this analysis, the Didymocyrtis clade is strongly
supported (ML-bs=72, PP=100) and includes species previ-
ously placed in Diederichia,Leptosphaeria, Phoma (or the
ne w genus Di e deric h omyce s recently desc ribed by
Tr ak un yingcharoen et al. 2014) and Polycoccum.
Didymocyrtis cladoniicola does not form a monophyletic
group, unlike the other species in the clade, which are strongly
supported by both analyses. However, relationships among
the speci e s a r e usually p o o r ly suppo r t e d. In the
Didymocyrtis consimilis clade, samples from Caloplaca hosts
cluster together, but this relationship is only supported by the
Bayesian analysis (PP=97).
Most taxa now treated in Didymocyrtis were previously
known either as species of Leptosphaeria or Polycoccum (sex-
ual stage) or of Diederichia or Phoma (asexual stage). Co-
occurrences of both stages on the same host thallus have rarely
been observed and documented until recently. Nevertheless,
our phylogenetic analyses clearly show that both stages be-
long to the same clade, representing the genus Didymocyrtis
(Figs. 1and 2), for which the much younger Diederichia and
Diederichomyces are made synonyms.
Fungal Diversity
Table 1 Specimens and DNA sequences used in this study, with their respective voucher information
Name Order Family Voucher information Hosts nuLSU ITS
Abrothallus parmotrematis Abrothallales Abrothallaceae AB1 KF816231
Acanthostigma minutum Tubeufiales Tubeufiaceae BBB:MVB 781 JN127360
Acrospermum compressum Acrospermales Acrospermaceae Vesterholt s.n. (TUR) EU940084
Anisomeridium ubianum Monoblastiales Monoblastiaceae Lumbsch; isolate 94 GU327709
Aptrootia robusta Trypetheliales Trypetheliaceae Lumbsch 20012 (F) KM453755
Architrypethelium uberinum Trypetheliales Trypetheliaceae Nelsen s.n. (F) KM453758
Arthonia didyma Arthoniales Arthoniaceae Ertz 7587 (BR) EU704083
Arthopyrenia cinchonae Trypetheliales Trypetheliaceae Lücking s.n. (F) KM453759
Arthopyrenia cf. planorbis Trypetheliales Trypetheliaceae Lücking 29584 JN872352
Arthopyrenia salicis Pleosporales Arthopyreniaceae CBS 368.94 AY538339
Asterina weinmanniae Asterinales Asterinaceae Hofmann 592 (M-0141091, PMA) GU586218
Astrothelium variolosum Trypetheliales Trypetheliaceae Nelsen s.n. (F) KM453768
Bathelium tuberculosum Trypetheliales Trypetheliaceae Lumbsch 19739z (F) KM453777
Bimuria novae-zelandiae Pleosporales Didymosphaeriaceae CBS 107.79 AY016356
Botryosphaeria dothidea Botryosphaeriales Botryosphaeriaceae CBS 115476 DQ678051
Caliciopsis pinea Coryneliales Coryneliaceae AFTOL-ID 1869 DQ678097
Capnodium coffeae Capnodiales Capnodiaceae CBS 147.52 DQ247800
Capronia munkii Chaetothyriales Herpotrichiellaceae AFTOL-ID 656 EF413604
Clypeococcum placopsiiphilum Trypetheliales Polycoccaceae Diederich 17576 (hb. Diederich) Placopsis gelida KT383789
Clypeococcum psoromatis Trypetheliales Polycoccaceae Ertz 19259 (BR) Squamarina cartilaginea KT383790
Coniosporium apollinis Unknown Unknown CBS 109860 GU250899
Coniothyrium telephii Pleosporales Coniothyriaceae CBS 188.71 GQ387599
Corynespora cassiicola Pleosporales Corynesporascaceae CBS 100822 GU301808
Cryomyces antarcticus Unknown Unknown CCFEE 536 GU250365
Cryptothelium pulchrum Trypetheliales Trypetheliaceae Nelsen 4001a (F) GU327714
Dacampia engeliana Pleosporales Dacampiaceae Hafellner 72868 (BR) Solorina saccata KT383791
Dacampia hookeri Pleosporales Dacampiaceae Hafellner 73897 (GZU) KT383792
Dacampia hookeri Pleosporales Dacampiaceae Hafellner 74269 (GZU) KT383793
Dacampia hookeri Pleosporales Dacampiaceae Hafellner 75980 (GZU) KT383794
Dacampia hookeri Pleosporales Dacampiaceae Hafellner 81840 (GZU) KT383795
Delitschia winteri Pleosporales Delitschiaceae CBS 225.62 DQ678077
Dendrographa leucophaea Arthoniales Roccellaceae Sparrius 7999 (BR) AY548810
Didymella bryoniae Pleosporales Didymellaceae CBS 133.96 GU456335
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Ertz 15238 (BR) Squamarina cartilaginea JQ238625 JQ238623
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Ertz 15208 (BR) Squamarina cartilaginea JQ238620
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Diederich 16915 (hb. Diederich) Cladonia sp. JQ238626
Fungal Diversity
Table 1 (continued)
Name Order Family Voucher information Hosts nuLSU ITS
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Diederich 16924 (hb. Diederich) Parmelina tiliacea JQ238629
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Van den Broeck 4662 (BR) Cladonia sp. JQ318014
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Ertz 16072 (BR) Ramalina pollinaria JQ318017
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Ertz 16069 (BR) Cladonia symphycarpa JQ318020
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Ertz 16064 (BR) Cladonia rangiformis JQ318023
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Ertz 16066 (BR) Cladonia foliacea JQ318026
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Ertz 16296 (BR) Ramalina polymorpha KT383809
Didymocyrtis cladoniicola Pleosporales Phaeosphaeriaceae Ertz 16464 (BR) Cladonia foliacea KT383810
Didymocyrtis consimilis Pleosporales Phaeosphaeriaceae Gardiennet 12041 –asexual stage Caloplaca cerina KT383812
Didymocyrtis consimilis Pleosporales Phaeosphaeriaceae Gardiennet 12041 –sexual stage Caloplaca cerina KT383796 KT383813
Didymocyrtis consimilis Pleosporales Phaeosphaeriaceae Freebury 1324 –asexual stage Caloplaca cerina JQ238635
Didymocyrtis consimilis Pleosporales Phaeosphaeriaceae Freebury 1357 –asexual stage Caloplaca cerina JQ238641
Didymocyrtis aff. consimilis Pleosporales Phaeosphaeriaceae Ertz 17617B (BR) –sexual stage Cladonia cf. pocillum KT383797 KT383811
Didymocyrtis aff. consimilis Pleosporales Phaeosphaeriaceae Berger 26876 –sexual stage Melanohalea exasperatula KT383798 KT383814
Didymocyrtis aff. consimilis Pleosporales Phaeosphaeriaceae Berger 27218 –asexual stage Melanohalea exasperatula KT383815
Didymocyrtis aff. consimilis Pleosporales Phaeosphaeriaceae Berger 27251 –asexual stage Heterodermia KT383816
Didymocyrtis aff. consimilis Pleosporales Phaeosphaeriaceae Diederich 17465 –asexual stage Cladonia pocillum KT383817
Didymocyrtis epiphyscia s. lat. Pleosporales Phaeosphaeriaceae Ertz 17422 (BR) Xanthoria parietina KT383818
Didymocyrtis epiphyscia s. lat. Pleosporales Phaeosphaeriaceae Ertz 17615 (BR) Physcia cf. adscendens KT383819
Didymocyrtis epiphyscia s. lat. Pleosporales Phaeosphaeriaceae Ertz 17411 (BR) Physcia adscendens KT383799 KT383820
Didymocyrtis epiphyscia s. lat. Pleosporales Phaeosphaeriaceae Ertz 17414 (BR) Xanthoria parietina KT383821
Didymocyrtis epiphyscia s. lat. Pleosporales Phaeosphaeriaceae Gardiennet 12008-isolate2 (BR) Xanthoria parietina KT383822
Didymocyrtis epiphyscia s. lat. Pleosporales Phaeosphaeriaceae Ertz 17461 (BR) Physcia adscendens KT383823
Didymocyrtis epiphyscia s. str. Pleosporales Phaeosphaeriaceae Freebury 1411; isolate 512 Physcia aipolia KT383824
Didymocyrtis epiphyscia s. lat. Pleosporales Phaeosphaeriaceae Ertz 17413 (BR) Xanthoria parietina KT383825
Didymocyrtis foliaceiphila Pleosporales Phaeosphaeriaceae Ertz 15257 (BR) Parmelia sulcata JQ318002
Didymocyrtis foliaceiphila Pleosporales Phaeosphaeriaceae Ertz 15258 (BR) Parmelia sulcata JQ318007 JQ318005
Didymocyrtis foliaceiphila Pleosporales Phaeosphaeriaceae Ertz 15262 (BR) Cladonia sp. JQ318008
Didymocyrtis foliaceiphila Pleosporales Phaeosphaeriaceae Ertz 15262bis (BR) Cladonia sp. JQ318011
Didymocyrtis foliaceiphila Pleosporales Phaeosphaeriaceae Diederich 17108 (hb. Diederich) Cladonia squamosa JQ238638
Didymocyrtis melanelixiae Pleosporales Phaeosphaeriaceae Harris 57475; isolate 551 –sexual stage Punctelia rudecta KT383826
Didymocyrtis melanelixiae Pleosporales Phaeosphaeriaceae Harris 57475; isolate 552 –sexual stage Punctelia rudecta KT383800 KT383827
Didymocyrtis melanelixiae Pleosporales Phaeosphaeriaceae Harris 57475; isolate 553 –asexual stage Punctelia rudecta KT383828
Didymocyrtis melanelixiae Pleosporales Phaeosphaeriaceae Harris 57475B; isolate 555 –asexual stage Punctelia rudecta KT383829
Didymocyrtis melanelixiae Pleosporales Phaeosphaeriaceae Harris 57465; isolate 557 –asexual stage Cetrelia olivetorum KT383830
Didymocyrtis melanelixiae Pleosporales Phaeosphaeriaceae Harris 57476; isolate 549 –asexual stage Punctelia rudecta KT383831
Fungal Diversity
Table 1 (continued)
Name Order Family Voucher information Hosts nuLSU ITS
Didymocyrtis pseudeverniae Pleosporales Phaeosphaeriaceae Diederich 17327a (hb. Diederich) Pseudevernia furfuracea KT383832
Didymocyrtis pseudeverniae Pleosporales Phaeosphaeriaceae Diederich 17327b (hb. Diederich) Pseudevernia furfuracea KT383833
Didymocyrtis pseudeverniae Pleosporales Phaeosphaeriaceae Diederich 17338 (hb. Diederich) Pseudevernia furfuracea KT383801 KT383834
Didymocyrtis ramalinae Pleosporales Phaeosphaeriaceae Paul 13i2013 isolate1 –sexual stage Ramalina fastigiata KT383835
Didymocyrtis ramalinae Pleosporales Phaeosphaeriaceae Paul 27i2013 isolate1 –sexual stage Ramalina fastigiata KT383836
Didymocyrtis ramalinae Pleosporales Phaeosphaeriaceae Paul 27i2013 isolate2 –sexual stage Ramalina fastigiata KT383837
Didymocyrtis ramalinae Pleosporales Phaeosphaeriaceae Ertz 16399 (BR) –sexual stage Ramalina sp. KT383802 KT383838
Didymocyrtis ramalinae Pleosporales Phaeosphaeriaceae Paul 10i2013 –asexual stage Ramalina fastigiata KT383839
Didymocyrtis ramalinae Pleosporales Phaeosphaeriaceae Van den Broeck 2900 (BR) –asexual stage Ramalina fastigiata JQ238614
Didymocyrtis slaptoniensis Pleosporales Phaeosphaeriaceae Gardiennet 12009 –sexual stage Xanthoria parietina KT383840
Didymocyrtis slaptoniensis Pleosporales Phaeosphaeriaceae MoraA –sexual stage Xanthoria parietina KT383803 KT383841
Didymocyrtis slaptoniensis Pleosporales Phaeosphaeriaceae MoraB –sexual stage Xanthoria parietina KT383842
Didymocyrtis xanthomendozae Pleosporales Phaeosphaeriaceae CBS129666 Xanthomendoza hasseana JQ238634 JQ238632
Dothidea insculpta Dothideales Dothideaceae CBS 189.58 DQ247802
Dothideomycetes sp. Unknown Unknown AN13 GU250928
Encephalographa elisae Eremithallales Melaspileaceae EB 0347 GU397343
Etayoa trypethelii Lichenostigmatales Phaeococcomycetaceae Common 9200-G (MSC, hb. Diederich) KF176940
Friedmanniomyces endolithicus Capnodiales Teratosphaeriaceae CCFEE 5180 GU250367
Geastrumia polystigmatis Asterinales? Unknown NC41.8F1a KF896877
Gloniopsis praelonga Hysteriales Hysteriaceae CBS 112415 FJ161173
Glonium circumserpens Mytilinidiales Gloniaceae CBS 123343; EB 0332 FJ161200
Halojulella avicenniae Pleosporales Halojulellaceae BCC 18422 GU371823
Hemigrapha atlantica Asterinales Parmulariaceae? Ertz 14014 (BR) KP456151
Hysterium angustatum Hysteriales Hysteriaceae CBS 236.34 FJ161180
Hysteropatella clavispora Patellariales Patellariaceae CBS 247.34 AY541493
Jahnula aquatica Jahnulales Aliquandostipitaceae R68-1 EF175655
Julella fallaciosa Trypetheliales Trypetheliaceae Nelsen s.n. (F) JN887400
Kellermania nolinae Botryosphaeriales Planistromellaceae CBS 131717 JX444876
Lachnum virgineum Helotiales Hyaloscyphaceae AFTOL-ID 49 AY544646
Laurera megasperma Trypetheliales Trypetheliaceae Ertz 9725 (BR) FJ267702
Lepidosphaeria nicotiae Pleosporales Testudinaceae CBS 101341 DQ678067
Leptosphaeria doliolum Pleosporales Leptosphaeriaceae CBS 125979 JF740283
Leptosphaeria macrospora Pleosporales Leptosphaeriaceae Kruys 501 (UPS) DQ384092
Lichenothelia convexa Lichenotheliales Lichenotheliaceae Diederich 17491 (hb. Diederich) KF176962
Lindgomyces ingoldianus Pleosporales Lindgomycetaceae ILLS:A-39-1B AB521736
Lophium mytilinum Mytilinidiales Mytilinidiaceae CBS 269.34 DQ678081
Fungal Diversity
Table 1 (continued)
Name Order Family Voucher information Hosts nuLSU ITS
Marcelaria purpurina Trypetheliales Trypetheliaceae Caceres 2009 KM453790
Massaria inquinans Pleosporales Massariaceae WU 30527 HQ599402
Massarina eburnea Pleosporales Massarinaceae CBS 473.64 GU301840
Melaspilea enteroleuca Eremithallales Melaspileaceae Aptroot 20685 (BR) KP456160
Montagnula opulenta Pleosporales Didymosphaeriaceae CBS 168.34 NG027581
Munkovalsaria appendiculata Pleosporales Didymosphaeriaceae CBS 109027 AY772016
Munkovalsaria donacina Pleosporales Didymosphaeriaceae Pitt s.n.; HVVV01 KJ628377
Mycomicrothelia hemispherica Trypetheliales Trypetheliaceae Lücking 28641 (F) GU327719
Mycomicrothelia miculiformis Trypetheliales Trypetheliaceae Lücking 28637 (F) GU327720
Mycomicrothelia obovata Trypetheliales Trypetheliaceae Nelsen 4007a (F) GU327721
Mycosphaerella punctiformis Capnodiales Mycosphaerellaceae CBS 113265 DQ470968
Myriangium duriaei Myriangiales Myriangiaceae CBS 260.36 DQ678059
Myrmaecium rubrum Valsariales Valsariaceae CBS 109505 GU456324
Neostagonospora caricis Pleosporales Phaeosphaeriaceae CBS 135092 KF251163
Neostagonospora elegiae Pleosporales Phaeosphaeriaceae CBS 135101 KF251164
Paraleptosphaeria orobanches Pleosporales Leptosphaeriaceae CBS 101.638 JF740299
Paraphoma radicina Pleosporales Phaeosphaeriaceae CBS 111.79 EU754191
Parastagonospora caricis Pleosporales Phaeosphaeriaceae Quaedvlieg S615 KF251176
Parastagonospora poae Pleosporales Phaeosphaeriaceae CBS 135091 KF251179
Phaeosphaeria lycopodina Pleosporales Phaeosphaeriaceae WA0000019137 JX981471
Phaeosphaeria oryzae Pleosporales Phaeosphaeriaceae CBS110110 GQ387591
Phaeotrichum benjaminii Phaeotrichales Phaeotrichaceae CBS 541.72 AY004340
Phyllobathelium anomalum Strigulales Strigulaceae Lücking s.n. (F) GU327722
Pleospora fallens Pleosporales Pleosporaceae CBS 284.70 GU238078
Pleospora herbarum Pleosporales Pleosporaceae CBS 191.86 GU238160
Polycoccum clauzadei Trypetheliales Polycoccaceae Freebury 2089 Xanthoria elegans KT383804
Polycoccum pulvinatum Trypetheliales Polycoccaceae Diederich 17389 (hb. Diederich) Physcia caesia KT383805
Polycoccum pulvinatum Trypetheliales Polycoccaceae Ertz 18114 (BR) Physcia caesia KT383806
Polycoccum trypethelioides Trypetheliales Polycoccaceae Diederich 17508 (hb. Diederich) Stereocaulon KT383807
Polycoccum vermicularium Trypetheliales Polycoccaceae Diederich 17545 (hb. Diederich) Thamnolia vermicularis KT383808
Polymeridium albocinereum Trypetheliales Trypetheliaceae Lücking s.n. (F) KM453795
Preussia terricola Pleosporales Sporormiaceae DAOM 230091 (AFTOL-ID 282) AY544686
Pseudopyrenula diluta Trypetheliales Trypetheliaceae Lücking 26062 (F) KM453797
Pseudotetraploa javanica Pleosporales Tetraplosphaeriaceae HHUF: 28596; Tanaka et al. AB524611
Pyrenochaeta nobilis Pleosporales Cucurbitariaceae CBS 292.74 GQ387615
Pyrenochaetopsis decipiens Pleosporales Cucurbitariaceae CBS 343.85 GQ387624
Fungal Diversity
Interestingly, our ITS phylogenetic tree highlights several
potential anamorph-teleomorph relationships, as follows: 1.
Phoma ficuzzae is shown to represent the asexual stage of
Didymocyrtis (Leptosphaeria)ramalinae; moreover, an asex-
ual Phoma-like stage (with pycnidia and conidia) was obtain-
ed in all our cultures of single ascospore isolates of
D. ramalinae. 2. ITS sequences of the asexual Phoma-like
stage and the sexual stage of Didymocyrtis melanelixiae are
identical and both stages clearly represent the same species. 3.
ITS sequences of the asexual Phoma caloplacae and the sex-
ual Didymocyrtis consimilis on Caloplaca gr. cerina are iden-
tical and both stages clearly represent the same species. 4. An
asexual Phoma-like stage (with pycnidia and conidia) was
obtained in our cultures of single ascospore isolates of
Didymocyrtis aff. consimilis (specimen Ertz 17617b, on
Cladonia cf. pocillum, initially identified as Polycoccum
laursenii), and the corresponding ITS sequence is identical
to the one of a Phoma-like anamorph collected in the same
locality and on the same host, C. pocillum (specimen
Diederich 17465).
In several other species both sexual and asexual stages some-
times occur on the same host thallus or apothecia, a phenomenon
that leads to the following hypotheses: 1. Phoma denigricans is
the asexual stage of D. bryonthae (both confined to Lecanora
epibryon). 2. An unnamed Phoma-like fungus occasionally ac-
companying perithecia of D. slaptoniensis on Xanthoria
parietina may be the asexual stage of that species. 3. Finally, a
sexual stage typical of Didymocyrtis has been found growing
with Phoma xanthomendozae, wherein the perithecia are inti-
mately intermixed with pycnidia and macroscopically indistin-
guishable from them, and is considered to represent the same
species.
Taxonomy
Pleosporales Luttr. ex M. E. Barr
Prodromus to class Loculoascomycetes (Amherst) 67
(1987). Type: Pleospora Rabenh. ex Ces. & De Not.
MycoBank: MB 90563.
Note. Pleosporales is the largest order of Dothideomycetes
with 41 families and includes saprobic fungi on dead plant
material, pathogens on living plants, hyperparasites on fungi
or insects, and lichenized species (Zhang et al. 2011; Hyde
et al. 2013).
Dacampiaceae Körb.
(as‘Dacampieae’), Syst. Lich. Germ.: 322 (1855). Type:
Dacampia A. Massal. MycoBank: MB 80680.
Notes. 1. Dacampiaceae was considered as a heteroge-
neous family in the past, with ten genera accepted by Hyde
et al. (2013). Our molecular data resolved the family as poly-
phylet i c . T h e g e n u s P o l y c o c c u m and members of
Clyp eococcum are accommodated in the new family
Polycoccaceae, while the genus Munkovalsaria is placed in
Table 1 (continued)
Name Order Family Voucher information Hosts nuLSU ITS
Racodium rupestre Capnodiales Unknown L346 - Hafellner & Muggia (GZU) EU048583
Setomelanomma holmii Pleosporales Phaeosphaeriaceae CBS 110217 GQ387633
Stagonospora foliicola Pleosporales Massarinaceae CBS 110111 KF251256
Stictographa lentiginosa Asterinales Unknown Ertz 17447 (BR) KP456169
Sympoventuria capensis Venturiales Sympoventuriaceae CBS-H 19757 DQ885904
Trypethelium nitidiusculum Trypetheliales Trypetheliaceae Ertz 9716 (BR) FJ267701
Trypethelium tropicum Trypetheliales Trypetheliaceae Nelsen s.n. (F) KM453819
Tubeufia cerea Unknown Tubeufiaceae CBS 254.75 DQ470982
Tyrannosorus pinicola Unknown Unknown CBS 124.88 DQ470974
Valsaria insitiva Valsariales Valsariaceae CBS 123125 GU460205
Venturia chlorospora Venturiales Venturiaceae Kruys 502 (UPS) DQ384101
Xanthoriicola physciae Capnodiales Teratosphaeriaceae Diederich 16713 (hb. Diederich) KF176965
GenBank accession numbers in bold refer to sequences (54) generated by this project. All other sequences (113 GenBank identification numbers) were obtained directly from GenBank
Fungal Diversity
Didymosphaeriaceae (Fig. 1). No molecular data are available
for the other genera currently placed in Dacampiaceae, i.e.
Aaosphaeria,Eopyrenula,Leptocucurthis,Pseudonitschkia,
Pyrenidium and Weddellomyces.
0.04
Racodium rupestre
Caliciopsis pinea
Bimuria novae-zelandiae
Glonium circumserpens
Dacampia hookeri 73897
Asterina weinmanniae
Didymocyrtis slaptoniensis MoraA [Polycoccum]
Arthopyrenia cinchonae
Polycoccum pulvinatum 17389
Setomelanomma holmii
Marcelaria purpurina
Hysteropatella clavispora
Pseudotetraploa javanica
Arthopyrenia cf. planorbis
Mycosphaerella punctiformis
Xanthoriicola physciae
Capnodium coffeae
Didymocyrtis foliaceiphila [Phoma]
Tyrannosorus pinicola
Polycoccum trypethelioides 17508
Cryomyces antarcticus
Stictographa lentiginosa
Dacampia hookeri 74269
Phaeosphaeria oryzae
Venturia chlorospora
Paraleptosphaeria orobanches
Sympoventuria capensis
Valsaria insitiva
Didymocyrtis ramalinae 16399 [Leptosphaeria]
Friedmanniomyces endolithicus
Capronia munkii
Pyrenochaetopsis decipiens
Lachnum virgineum
Didymocyrtis xanthomendozae [Phoma]
Corynespora cassiicola
Massaria inquinans
Mycomicrothelia hemispherica
Polycoccum vermicularium 17545
Myriangium duriaei
Etayoa trypethelii
Cryptothelium pulchrum
Didymocyrtis aff. consimilis 26876 [Polycoccum]
Anisomeridium ubianum
Didymocyrtis pseudeverniae 17338 [Diederichia]
Paraphoma radicina
Clypeococcum placopsiiphilum 17576
Didymella bryoniae
Massarina eburnea
Laurera megasperma
Halojulella avicenniae
Leptosphaeria doliolum
Melaspilea enteroleuca
Mycomicrothelia miculiformis
Leptosphaeria macrospora
Preussia terricola
Clypeococcum psoromatis 19259
Didymocyrtis epiphyscia s. lat. 17411 [Phoma]
Pleospora fallens
Astrothelium variolosum
Phaeotrichum benjaminii
Acanthostigma minutum
Munkovalsaria appendiculata
Lichenothelia convexa
Tubeufia cerea
Coniothyrium telephii
Pyrenochaeta nobilis
Arthonia didyma
Architrypethelium uberinum
Lepidosphaeria nicotiae
Dendrographa leucophaea
Phyllobathelium anomalum
Pleospora herbarum
Delitschia winteri
Aptrootia robusta
Botryosphaeria dothidea
Polymeridium albocinereum
Dacampia engeliana 72868
Jahnula aquatica
Munkovalsaria donacina
Didymocyrtis cladoniicola [Phoma]
Gloniopsis praelonga
Didymocyrtis melanelixiae 57475 [Leptosphaeria]
Geastrumia polystigmatis
Bathelium tuberculosum
Didymocyrtis consimilis s. str. 12041 [Polycoccum]
Dothideomycetes AN13
Dacampia hookeri 75980
Lophium mytilinum
Julella fallaciosa
Pseudopyrenula diluta
Dacampia hookeri 81840
Polycoccum pulvinatum 18114
Montagnula opulenta
Trypethelium nitidiusculum
Kellermania nolinae
Mycomicrothelia obovata
Arthopyrenia salicis
Hysterium angustatum
Myrmaecium rubrum
Didymocyrtis aff. consimilis 17617B [Polycoccum]
Dothidea insculpta
Coniosporium apollinis
Lindgomyces ingoldianus
Acrospermum compressum
Abrothallus parmotrematis
Polycoccum clauzadei 2089
Encephalographa elisae
Hemigrapha atlantica
Trypethelium tropicum
POLYCOCCACEAE
fam. nov.
PHAEOSPHAERIACEAE
PLEOSPORALES
TRYPETHELIALES
100
100
95
100
100
100
100
99
100
100
100
100
100
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100
99 100
97
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100
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99
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100
DACAMPIACEAE
DIDYMOSPHAERIACEAE
TRYPETHELIACEAE
Fig. 1 Phylogenetic relationships among 109 samples within
Dothideomyceta (with three outgroup taxa) based on a data set of
nuLSU sequences that resulted from a Maximum Likelihood analysis
using Garli. Internal branches with Maximum Likelihood bootstrap
values ≥70 obtained from a Garli analysis are considered strongly
supported and represented by thicker lines. Posterior probabilities ≥95
resulting from a Bayesian analysis are shown above internal branches.
The newly sequenced specimens are in bold. The old and most commonly
used generic name is indicated within brackets for the species now placed
in Didymocyrtis. Collecting numbers of the authors following the species
names act as specimen and sequence identifiers
Fungal Diversity
2. The genus Da campi a is related to members of
Leptosphaeriaceae in our nuLSU tree (e.g., Paraleptosphaeria
orobanches, Fig. 1). The relationship of Dacampiaceae with
other families of Pleosporales will require more detailed
study. Leptosphaeriaceae will become a synonym of
Dacampiaceae if Dacampia is shown to be nested with-
in it.
Dacampia A. Massal. (Fig. 3)
Sulla Lec. Hook. Schaer.: 7 (1853). Type: Dacampia
hookeri (Borrer) A. Massal. MycoBank: MB 1401.
0.02
D. slaptoniensis 12009, France/Xanthoria parietina [Polycoccum]
D. melanelixiae 57475B-555, USA/Punctelia rudecta [Phoma]
D. foliaceiphila JQ238638, Belgium/Cladonia squamosa [Phoma]
D. cladoniicola JQ238623, Belgium/Squamarina cartilaginea [Phoma]
D. aff. consimilis 27251, Madeira/Heterodermia [Phoma]
D. foliaceiphila JQ318011, Belgium/Cladonia sp. [Phoma]
D. pseudeverniae 17338, Switzerland/Pseudevernia furfuracea [Diederichia]
Neostagonospora elegiae
D. consimilis JQ238635, Canada/Caloplaca cerina [Phoma]
D. epiphyscia s. str. 1411, Canada/Physcia aipolia [Phoma]
D. epiphyscia s. lat. 17461, France/Physcia adscendens [Phoma]
D. slaptoniensis MoraA, France/Xanthoria parietina [Polycoccum]
D. ramalinae HP10i2013-is1, Scotland/Ramalina fastigiata [Phoma]
D. consimilis 12041, France/Caloplaca cerina [Polycoccum]
D. epiphyscia s. lat. 17413, Belgium/Xanthoria parietina [Phoma]
D. epiphyscia s. lat. 12008-is2, France/Xanthoria parietina [Phoma]
D. melanelixiae 57475-553, USA/Punctelia rudecta [Phoma]
D. cladoniicola JQ318014, Belgium/Cladonia sp. [Phoma]
D. cladoniicola JQ318020, France/Cladonia symphycarpa [Phoma]
D. aff. consimilis 26876, Austria/Melanohalea exasperatula [Polycoccum]
D. melanelixiae 57476-549, USA/Punctelia rudecta [Phoma]
D. ramalinae JQ238614, France/Ramalina fastigiata [Phoma]
D. xanthomendozae JQ238632, Canada/Xanthomendoza hasseana [Phoma]
D. pseudeverniae 17327a, Switzerland/Pseudevernia furfuracea [Diederichia]
D. cladoniicola JQ238620, Belgium/Squamarina cartilaginea [Phoma]
D. ramalinae 16399, Corsica/Ramalina [Leptosphaeria]
Neostagonospora caricis
D. aff. consimilis 17617b, Luxembourg/Cladonia cf. pocillum [Polycoccum]
Phaeosphaeria lycopodina
D. slaptoniensis MoraB, France/Xanthoria parietina [Polycoccum]
D. melanelixiae 57475-551, USA/Punctelia rudecta [Leptosphaeria]
Stagonospora foliicola
D. melanelixiae 57475-552, USA/Punctelia rudecta [Leptosphaeria]
D. cladoniicola JQ318017, France/Ramalina pollinaria [Phoma]
D. consimilis 12041, France/Caloplaca cerina [Phoma]
D. epiphyscia s. lat. 17411, Belgium/Physcia adscendens [Phoma]
D. cladoniicola JQ238629, Mallorca/Parmelina tiliacea [Phoma]
D. epiphyscia s. lat. 17615, Luxembourg/ Physcia cf. adscendens [Phoma]
D. epiphyscia s. lat. 17414, Belgium/Xanthoria parietina [Phoma]
D. cladoniicola 16464, Corsica/Cladonia foliacea [Phoma]
D. aff. consimilis 17465, Luxembourg/Cladonia pocillum [Phoma]
D. cladoniicola JQ238626, Mallorca/Cladonia sp. [Phoma]
Parastagonospora poae
D. foliaceiphila JQ318005, Belgium/Parmelia sulcata [Phoma]
D. foliaceiphila JQ318002, Belgium/Parmelia sulcata [Phoma]
D. epiphyscia s. lat. 17422, Belgium/Xanthoria parietina [Phoma]
D. aff. consimilis 27218, Austria/Melanohalea exasperatula [Phoma]
D. ramalinae HP13i2013-is1, Scotland/Ramalina fastigiata [Leptosphaeria]
D. ramalinae HP27i2013-is2, Scotland/Ramalina fastigiata [Leptosphaeria]
D. cladoniicola JQ318026, France/Cladonia foliacea [Phoma]
D. consimilis JQ238641, Canada/Caloplaca cerina [Phoma]
D. pseudeverniae 17327b, Switzerland/Pseudevernia furfuracea [Diederichia]
D. cladoniicola JQ318023, France/Cladonia rangiformis [Phoma]
Parastagonospora caricis
D. foliaceiphila JQ318008, Belgium/Cladonia sp. [Phoma]
D. cladoniicola 16296, Canary Islands/Ramalina polymorpha [Phoma]
D. ramalinae HP27i2013-is1, Scotland/Ramalina fastigiata [Leptosphaeria]
D. melanelixiae 57465-557, USA/Cetrelia olivetorum [Phoma]
100
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DIDYMOCYRTIS
Fig. 2 Phylogenetic relationships among 52 samples of Didymocyrtis
based on a data set of ITS sequences that resulted from a Maximum
Likelihood analysis using Garli. Internal branches with Maximum
Likelihood bootstrap values ≥70 obtained from a Garli analysis are
considered strongly supported and represented by thi cke r lines.
Posterior probabilities ≥95 resulting from a Bayesian analysis are
shown above internal branches. The newly sequenced specimens are in
bold. Collecting numbers of the authors following the species names act
as specimen and sequence identifiers. The geographical origin and the
host lichen species of the sequenced specimens of Didymocyrtis are also
indicated. The old and most commonly used generic name is indicated
within brackets for the species now placed in Didymocyrtis. The length of
the branch represented by dashed lines was reduced by 50 % for editing
reason
Fungal Diversity
Syn.: Xenosphaeria Trevis., Conspect. Verruc.: 18 (1860).
Type: Xenosphaeria hookeri (Schaer.) Trev., lectotypified by
Clements and Shear (1931). MycoBank: MB 5823.
Lichenicolous or lichenized with Coccomyxa and external
cephalodia with Nostoc. Ascomata perithecioid, with a central
ostiole, lacking setae, black, immersed to semi-immersed,
with a pseudoparenchymatic ascomatal wall, in longitudinal
section seen as a textura angularis with thick-walled, reddish
to dark brown cells. Hamathecium of interascal filaments
(paraphysoids) with ramifications and anastomoses. Asci
bitunicate, subcylindrical, apically thickened when mature,
with a small, sometimes indistinct, ocular chamber, 8-spored,
ascal wall and hymenial gel I–, more rarely K/I+ bluish.
Ascospores 1–2-seriate, pale brown to dark brown,pigmented
from an early stage of development, muriform, ellipsoid to
fusiform, often constricted at the septa, smooth, without visi-
ble gelatinous sheath. Asexual stage unknown.
Note. The genus Dacampia includes 15 species, most
of them being lichenicolous. The type species,
Dacampia hookeri, is lichenized (except juvenile stages
that might facultatively transform the thallus of Solorina
bispora), while the lichenicolous Dacampia engeliana
modifies its host lichen to form a thallus structure sim-
ilar to that found in D. hookeri (Henssen 1995; Hyde
et al. 2013; de los Ríos and Grube 2000). The genus
might be heterogeneous, as so m e l i c h e n i colous
Dacampia species that do not form such thallus struc-
tures might be more distantly related, but molecular data
are currently missing for them.
Phaeosphaeriaceae M. E. Barr
Mycologia 71: 948 (1979). Type: Phaeosphaeria I.
Miyake. MycoBank: MB 81637.
Note. Phaeosphaeriaceae is a large and important family in
the Pleosporales (Hyde et al. 2013, Phookamsak et al. 2014).
Lawrey et al. (2012) placed all lichenicolous Phoma species in
Phaeosphaeriaceae, while de Gruyter et al. (2009,2013) stat-
ed that Phoma s . str. s hould be restr icted only to
Didymellaceae. Eventually, Trakunyingcharoen et al. (2014)
described the new genus Diederichomyces to accommodate
most of the lichenicolous Phoma. But as shown here, several
older generic names are available for the lichenicolous Phoma
species, of which Didymocyrtis is the oldest.
Fig. 3 Morphological features of the genus Dacampia.a,c–fDacampia
hookeri (athallus with marginal ascomata; chymenium; d–easci; f
ascospores; in water; Hafellner 75980). bD. engeliana on Solorina
(Germany, Bad Tölz, 1978, Feuerer). Scale bars:a–b=200 μm; c=
50 μm; d–f=5 μm
Fungal Diversity
Didymocyrtis Vain.
Acta Soc. Fauna Flora Fenn. 49(2): 221, 263 (1921). Type:
Didymocyrtis consimilis Vain., lectotype, selected here.
MycoBank: MB 1554.
Syn. nov.: Diederichia D. Hawksw., Lichenologist 35: 206
(2003). Type: Diederichia pseudeverniae (Etayo & Diederich)
D. Hawksw. MycoBank: MB 28744.
Syn. nov.: Diederichomyces Crous & Trakunyingcharoen in
Trakunyingcharoen et al., IMA Fungus 5: 393 (2014). Type:
Diederichomyces xanthomendozae (Diederich & Freebury)
Crous & Trakunyingcharoen. MycoBank: MB 810828.
As coma ta (unknown in some species) perithecioid
(pseudothecia); ascomatal wall dark brown, pseudoparenchyma-
tous, in longitudinal section consisting of polyhedral, tangentially
flattened cells forming a textura angularis, with the pigment de-
posited in the hyphal cell walls. Hamathecial filaments
paraphysoids, persistent, septate, with some branches and anas-
tomoses. Asci fissitunicate, narrowly cylindrical, endoascus lat-
erally thickened when young, apically thickened when mature,
with a small ocular chamber, ascal wall I–and K/I–except
dextrinoid reaction of ascal periplasma, hymenial ‘gel’I–and
and K/I–, 8-spored. Ascospores ± uniseriate to half-overlapping,
pale brown to brown, usually pigmented from an early stage of
development (mature ones pale brown only in some species),
relatively thin-walled (compared to those of true Polycoccum-
species), transversally 1–3-septate, upper half-spore slightly
broader than lower one, with rounded ends, most species with
a distinct sculpture in light microscopy, a distinct perispore visi-
ble in young spores of some species in K.
Conidiomata (unknown in some species) Phoma-like, with
unilocular, ostiolate pycnidia; pycnidia usually undistinguishable
from perithecia, except sometimes by the smaller size; pycnidial
wall similar to perithecial wall. Conidiophores lacking.
Conidiogenous cells attached to the conidiomal wall and lining
the cavity, hyaline; conidiogenesis phialidic, not proliferating.
Conidia hyaline, simple, smooth-walled, not embedded in a ge-
latinous matrix, usually with rounded ends.
Notes. 1. Boerema (1997) subdivided the genus Phoma
into nine sections with sexual stages in the genera
Didymella, Leptosphaeria, Mycosphaerella and Pleospora.
de Gruyter et al. (2009) confirmed the polyphyletic character
of Phoma in the Pleosporineae by the application of molecu-
lar methods (see also Zhang et al. 2009). The generic type,
Phoma herbarum, grouped in the Didymellaceae, and there-
fore, Phoma s. str. is considered to be restricted to the
Didymellaceae. Aveskamp et al. (2010) confirmed that
Phoma species appear not only in the Didymellaceae, but also
in the related clades Leptosphaeriaceae, Phaeophaeriaceae
and Pleosporaceae. In a recent phylogenetic study focusing
on species of Sphaerellopsis, Trakunyingcharoen et al. (2014)
described the new genus Diederichomyces as including most
of the lichenicolous Phoma species that had been assigned to
the Phaeosphaeriaceae by Lawrey et al. (2012). As shown
here, the genus Diederichia also belongs to this clade.
However, the generic name Didymocyrtis Vain., neglected
practically from its publication (Vainio 1921), is also available
and is here used for this group (see Taxonomy section). As a
consequence Diederichia and Diederichomyces are synony-
mized with Didymocyrtis.
2. Vainio (1921) described both the genus Didymocyrtis
and the species D. consimilis in detail, and mentioned
D. physciicola (Nyl.) Vain. (based on Mycoporum physciicola
Nyl., a later heterotypic synonym of Sphaerellothecium
parietinarium (Linds.) Hafellner & V. John) and Mycoporum
epistygium Nyl. (Vainio 1921: 222) (an ascomycete of unclear
relationship on Melanelia stygia). Mycoporum epistygium
was combined into Didymocyrtis epistygia (Nyl. ex Vain.)
Vain. later in the same publication (Vainio 1921: 263) and,
therefore, Didymocyrtis consimilis cannot be treated as the
holotype and is here selected as lectotype.
3. Didymocyrtis differs from Polycoccum s. str. mainly by
narrowly cylindrical asci, ± monostichously arranged asco-
spores, thin paraphysoids, thin-walled, medium (reddish)-
brown ascospores and a Phoma-like anamorphic stage. As
well, Polycoccum s. str. often induces galls formation on the
host lichens, whereas Didymocyrtis does not.
4. Many other lichenicolous Phoma-like species have been
described in the past (Lawrey and Diederich 2015), and some
of these will certainly be combined later in Didymocyrtis, as
soon as sequences of them become available. However, these
species should not be combined in this genus, based on mor-
phological characters alone. E.g., sequences of Phoma
puncteliae Diederich & Lawrey placed this species ouside of
Didymocyrtis (Lawrey et al. 2012) and Trakunyingcharoen
et al. (2014) introduced the new genus Xenophoma to accom-
modate the species. Similarly, Phoma cytospora (Vouaux) D.
Hawksw. has been shown to belong to Arthoniales, and the
new genus Briancoppinsia has subsequently been described
for it (Diederich et al. 2011).
5. Didymocyrtis is also a generic name used for protists in
the family Coccodiscidae (e.g. Gilg et al. 2010).
Key to species of Didymocyrtis
1 Fructifications are perithecioid ascomata.......................2
1 Fructifications are pycnidial conidiomata....................9
2 Ascosporespredominantly 2-or 3-septate....................3
2 Ascospores predominantly 1-septate............................4
3 Ascospores predominantly 2-septate, some 1- and 3-
sep t at e i n te r mi x e d, u p t o 15 μm l o ng; o n
Parmeliaceae.......................................D. melanelixiae
3 Ascospores predominantly 3-septate, some 1- or 2-
septate ascospores intermixed, usually longer than
15 μm; on Ramalina [specimens on Protousnea,
named Leptosphaeria protousneae, will also key out
here]..........................................................D. ramalinae
Fungal Diversity
4 Ascospores 16–2 0 × 8–10 μm; on Tel osc histes
......................................................................D. kaernefeltii
4 Ascospores up to 15×7 μm long......................................5
5 Many asci longer than 90 μm; ascospores 13–15 ×6–
7μm.............................................................................6
5 Most asci up to 90 μm long.........................................7
6 Asco s p ore s 1 3 –15 × 6 –7μm; on X a nt h o r i a
parietina..............................................D. slaptoniensis
6 A s c o s p o r e s 1 1 –1 3 × 6 –7μm ; o n
Xanthomendoza..........................D. xanthomendozae
7 All ascospores 1-septate, 10–13 ×4.5–6μm; on
Teloschistes................................................D. infestans
7 Some 2-septate ascospores intermixed; on apothecia of
crustose lichens.............................................................8
8 Verruculose sculpture of ascospores distinct; asco-
spores 11.5–14 ×4–5μm; on Lecanora epibryon
[coni d i a, if pre s e nt, ± el l i psoid , 7 –8 × 3 –
4μm].......................................................D. bryonthae
8 Sculpture of ascospores very minute and often indistinct;
ascospores 12–15×5–6μm; on Caloplaca [conidia, if
present, broadly ellipsoid, 4.5–6.5 × 2.5–4.5 μm, to
subglobose, 5–6μm diam.]...........................D. consimilis
9 Conidia mostly 16–22×6–9μm, multiguttulate; on
Pseudevernia furfuracea...................D. pseudeverniae
9 Conidia smaller, less than 8 × 6 μm, usually 1–2-
guttulate.....................................................................10
10 Conidia broadly ellipsoid, l/b ratio<1.6, mostly 1-
guttulate....................................................................11
10 Conidia narrowly ellipsoid, l/b ratio>1.4, mostly 2-
guttulate....................................................................13
11 Conidia mostly 3.8–5.1 ×3.2–3.8 μm, with one large
guttule; on Par meliace ae [some specimens of
D. consimilis s. lat. and of D. ramalinae may also
key out here] .......................................D. melanelixiae
11 Conidia at least 4.5×3.5 μm....................................12
12 Pycnidia (50–)100–150 μm diam.; conidia broadly
ellipsoid; on Physcia aipolia........D. epiphyscia s. str.
12 Pycnidia 70–100(–130) μm diam.; conidia broadly
ellipsoid to subspherical; on Caloplaca (gr. cerina or
gr. tiroliensis) [D. consimilis s. lat. also on Cladonia
po c i l lum ,Heterodermia and M e l an o h a le a
exasperatula]..........................................D. consimilis
13 Conidia very narrowly ellipsoid, l/b ratio mainly 2.4–
3; on Cladonia and Parmelia ..............D. foliaceiphila
13 Conidia narrowly ellipsoid, l/b ratio mainly 1.4–
2.3............................................................................14
14 Conidial breadth mainly<3 μm...............................15
14 Conidial breadth mainly>3 μm...............................17
15 Co nid i a 6–8 × 2 .5–3. 5 μm ; on X an t ho r i a
parietina..................................................D. slaptoniensis
15 Conidia shorter, mainly less than 6.5×3 μm [two ge-
netically distinct species that can hardly be distin-
guished morphologically].........................................16
16 Pycnidia (50–)100–150 μm diam.; on Phys cia
adscendens,P. tenella and Xanthoria parietina
.....................................................D. epiphyscia s. lat.
16 Pycnidia (40–)50–100(–140) μm; on Cladonia,
Ra m a l ina ,S q ua m a r in a an d P arm e l i ac e a e
...........................................................D. cladoniicola
17 Conidial l/b ratio > 2..................................................18
17 Conidial l/b ratio <2 [two genetically distinct species
of which the asexual stages are morphologically very
similar]......................................................................19
18 Conidia mainly 7–8×3–4μm; on Lecanora epibryon
(hymenium) [ascospores, if present, mainly 11.5–14×
4–5μm]..................................................D. bryonthae
18 Conidia mainly 6–8 × 2.5–3.5 μm; on Xanthoria
parietina [ascospores, if present, mainly 13–15×6–
7μm]..................................................D. slaptoniensis
19 Pycnidia 105–135 μm diam.; on Ramalina [asco-
spores, if present, 3-septate].....D. ramalinae
19 Pycnidia 14 0–160 μm; o n X antho mend o za
[ a s c o s p o r e s , i f p r e s e n t , 1 -
septate]..................................D. xanthomendozae
Didymocyrtis bryonthae (Arnold) Hafellner comb. nov.
Endococcus bryonthae Arnold, Flora (Regensburg) 57:
141 ([21. März] 1874); Didymosphaeria bryonthae (Arnold)
G. Winter, Rabenh. Krypt.-Fl., 2. Aufl., 1(2): 430 (1885);
Microthelia bryonthae (Arnold) Kuntze, Revisio generum
plantarum 3: 498 (1898); Mycoporum bryonthae (Arnold)
Jatta, Syll. Lich.: 494 (1900); Tichothecium bryonthae
(Arnold) Jatta, Fl. Ital. Crypt. Lich.: 841 (1911); Sphaeria
bryonthae (Arnold) H. Olivier, Bull. Int. Acad. Géogr. Bot.
17: 170 (1907); Polycoccum bryonthae (Arnold) Vězda,
Ceská Mycol. 23: 109 (1969). Type: [Austria, Nordtirol,
Serlos-Gruppe], Kalkboden links ober der Ochsenalm,
Matreier Grube, Waldrast in Tirol, 6000′[Wiener Fuß = c.
1900 m alt.], on Lecanora epibryon (as Lecanora subfusca
var. bryontha), Aug. 1873, F. Arnold (M –holotype!).
MycoBank: MB 814022
Syn.?: Phoma denigricans Hafellner, Herzogia 10: 18
(1994). Type: Italy: Trentino, Dolomiten, Pordoi-Joch, N-
Fuß des Sass Beccle, 46°29′05″N, 11°48′40″E, c. 2300 m,
Hänge mit niedrigen Dolomitschrofen, on Lecanora epibryon
(apoth.), 25 Oct. 1984, J. Hafellner 11989 (GZU –holotype!).
MycoBank: MB 447502
Ascomata in dense groups on the apothecia of the host,
almost completely immersed in the apothecial hymenium
that becomes black and might finally bleach, more pro-
tr uding whe n the host hy menium ha s collapsed,
subspherical, 150–200(–250) μm diam.; wall dark brown,
often somewhat paler towards the base, laterally c. 12–
15 μm diam., of about 4–6 layers of cells. Paraphysoids
1–1.5 μm diam. Asci 60–80×7–10 μm. Ascospores ±
uniseriate, pale brown, 1-septate, some (less than 5 %)
Fungal Diversity
with an additional septum in the upper cell, (10–)11.5–
14(–16)×4–5μm, often slightly constricted at the septum;
septum with externally protruding torus; distin ct
verruculose sculpture visible in light microscopy.
Conidiomata of supposed asexual stage immersed in the
hymenium of host apothecia, several per apothecium, black-
ening the apothecial surface, in longitudinal section broadly
oval to subspherical, c. 80–100×100–130 μm diam.; wall
dark brown above, paler brown towards the base, of c. 3–5
layers of cells; conidiogenous cells broadly pear-shaped, 7–
9× 4–5μm. Conidia ellipsoid to somewhat tapering towards
the lower end to oblong, many biguttulate, (6–)7–8×3–4μm,
l/b ratio 2–2.3.
For further descriptions and illustrations of the teleomorph
see Arnold (1874), and of the supposed anamorph see
Hafellner (1994).
Distribution and hosts. Taking into consideration pub-
lished records, and adding the data for the supposed
anamorph (Phoma denigricans) and of the specimens cited
below, the known distribution can be summarized as fol-
lows: Europe (Austria, Germany, Italy, Poland, Svalbard,
Sweden), Asia (Mongolia, Russia), and North America
(C anada, G reenla nd, USA) ; in the hymenium of
Lecanora epibryon (e.g., Arnold 1874; Hafellner 1994;
Kukwa and Flakus 2009; Santesson 1993; Triebel and
Scholz 2001; Zhurbenko 1996,2009b).
Notes. 1. The collection Arnold, Lich. exs. 615 sub
Endococcus bryonthae given as type by various authors
(e.g., Aptroot 1995b) is in fact a topotype; that specimen
was collected on 4 Sept. 1874 and therefore after the appear-
ance of the protologue (March 1874).
2. Hawksworth and Diederich (1988: 297) and Aptroot
(1995b: 59) erroneously described the host of Arnold, Lich.
exs. 615 (sec. Arnold duplicates of a later collection from the
locus classicus) as Pertusaria bryontha; however, according
to Arnold and the specimens seen by us, it is Lecanora
epibryon.
3. Sometimes ascomata of Didymocyrtis bryonthae
and pycnidia of Phoma denigricans can be observed
on the same specimen (e.g., Hafellner 38111, 76036,
including on one host apothecium). Comparable obser-
vations have also been reported from Svalbard and
Siberia by Zhurbenko (2009b: 147, 148). Such co-
occurrences are one of the arguments for our opinion
that the two taxa represent the sexual and asexual stages
of the same fungal species.
4. For differences between Didymocyrtis bryonthae and
D. consimilis, see notes below under D. consimilis.
5. The record on Lecanora bryopsora (Hafellner 2008)
could not be confirmed. The specimen, on which this record
was based, shows infested L. epibryon growing intermingled
with an undetermined sorediate crust which evidently is not
L. bryopsora.
Selected specimens examined (all on Le cano ra
e p i b r y o n ) : S e x u a l s t a g e: A u s t r i a : S a l z b u rg ,
Nationalpark Hohe Tauern, Glockner Gruppe, NW-Grat
des Großen Margrötzen Kopfs W ober dem Hochtor,
47°05′10″N, 12°50′10″E, 2620 m, GF 8943/1, 1996,
Hafellner 38111 & Wittmann (hb. Hafellner) (with asex-
ual stage). Steiermark, Nordalpen (Nördliche Kalkalpen),
Totes Gebirge, Hochangern-Massiv N von Liezen,
Nazogl, knapp NE vom Gipfel auf dem Rücken gegen
den Angerkogel, 47°36′45″N, 14°13′50″E, 2050 m,
GF 8351/3, 2010, Hafellner 76036 & Muggia (GZU)
(with asexual stage). Greenland: W-Grönland, Gem.
Umanak, Hänge über Marmorilik, 50–300 m, 1983,
Po elt & Ullrich ( GZU). Italy : Lombardi a, prov.
Brescia, Eastern Alps, Central Alps, Ortler-group
(Stelvio-group), Cima di Cadì N above Passo del
Tonale, N below the summit, 46°16′35″N, 10°34′15″
E, 2590 m, 2006, Hafellner 75406 & Muggia (GZU).
Mongolia: Chubsugul Chimak, Tomim-Brigade, 1983,
Huneck MVR 83-43 (GZU). Russia: Siberia, Yakutiya
(Sakha-Yakutiya Republic), Lena River delta, right bank
of Olenek channel/river branch, near fishermans’house
Novyi Chai-Tumus, 45 WNW Krest-Tumsa cape and
Sokol polar station, 72°22′N, 125°40′E, 10–40 m,
1998, Zhurbenko 9896 (M).
Asexual stage: Austria: Steiermark, Zentralalpen, Niedere
Tauern, Schladminger Tauern, SE-Abhänge der Steirischen
Kalkspitze, W der Giglachseehütte zwischen Preuneggsattel
und Akarscharte, 47°16′50″–17′00″N, 13°37′50″–38′10″E,
1970–2080 m, GF 8747/2, 2001, Obermayer 9166 (GZU).
Germany: Bayern, Landkreis Berchtesgadener Land,
Steinernes Meer, Viehkogel, auf dem Gipfel, 2150 m, GF
8543, 1985, Wunder 4030 & Türk (M). Greenland: W-
Grönland, Gem. Umanak, Marmorilik, N-seitige Hänge S
über dem Fjord Qaumarujuk, 5–200 m, 1983, Poelt &
Ullrich (GZU).
Didymocyrtis cladoniicola (Diederich, Kocourk. &
Etayo) Ertz & Diederich comb. nov.
Phoma cladoni icola Diederich, Kocourk. & Etayo,
Lichenologist 39: 157 (2007); Diederichomyces cladoniicola
(Diederich, Kocourk. & Etayo) Crous & Trakunyingcharoen
in Trakunyingcharoen et al., IMA Fungus 5: 401 (2014).
Type: USA, Minnesota, Cottonwood County, 2 miles W of
US 71, 14.5 miles N of Windom, 44°06′40″N, 95°04′30″
W, 400 m, seasonally dry, flat, Sioux quartzite outcrop on N
side of road, on Cladonia pyxidata, 28 Sept. 1991, W. R. Buck