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Persoonia 39, 2017: 91–117 ISSN (Online) 1878-9080
www.ingentaconnect.com/content/nhn/pimj https://doi.org/10.3767/persoonia.2017.39.05
RESEARCH ARTICLE
INTRODUCTION
Lichenicolous fungi are a group of fungi specialized in living on
lichens as parasites, commensals or saprotrophs (Hawksworth
2003, Lawrey & Diederich 2003). About 2 000 species of licheni-
colous fungi have been described, 96 % of them belonging to
the Ascomycota and the rest to the Basidiomycota (Lawrey
& Diederich 2016). However, it is assumed that their species
diversity is much greater (Hawksworth & Rossman 1997), the
estimated total number of species lying between 3 000 and
4 000 (Hawksworth 2001, Lawrey & Diederich 2003). From
approximately 15 % of the described species only the asexual
stage is known (Lawrey & Diederich 2016) and the taxonomical
affiliation of most of them is uncertain. The generic concepts of
asexual fungi are still based on morphological characters and
numerous changes are to be expected in the future. Just as
in fungi with other lifestyles, phylogenetic studies have proved
that many genera are polyphyletic (Verkley & Starink-Willemse
2004, Crous et al. 2007, Aveskamp et al. 2010, De Gruyter et al.
2010). The biology and ways of interaction of lichenicolous fungi
with their hosts are still rather poorly known, although some
anatomic studies have been carried out. Lichenicolous basidi-
omycetous fungi, most of which belong to the Tremellomycetes,
generally induce the formation of galls. Both the host and the
parasite hyphae take part in these galls, while the photobiont
does not intervene in their production (Grube & De los Ríos
2001). The interaction of ascomycetous lichenicolous fungi with
their hosts is more varied: some of them also induce galls, oth-
ers produce necrotic areas on the lichen thallus, and others do
not produce any morphological change in the thallus (Rambold
& Triebel 1992). As for the connections with the host, often the
lichenicolous fungus hyphae reach the algal layer, where they
form haustoria with the photobiont, while some species estab-
lish connections with the mycobiont (Rambold & Triebel 1992,
De los Ríos & Grube 2000, De los Ríos et al. 2000).
More than a decade ago DNA sequences began to be used in
order to determine the phylogenetic placement of lichenicolous
fungi (e.g., Peršoh & Rambold 2002, Hawksworth et al. 2010,
Ruibal et al. 2011, Suija et al. 2015), but this work has been
much slower than in other groups of fungi, essentially due to
the small size of most lichenicolous fungi, the risk of a con-
tamination with the host material and the difficulty of obtaining
axenic cultures. The lichenicolous lifestyle is present in seven
classes within the Ascomycota (Lawrey & Diederich 2016),
but their abundance is not the same in all of them (Arnold et
al. 2009). A high number of lichenicolous species belong to
Lecanoromycetes (Rambold & Triebel 1992, Lawrey & Diede
rich 2003, Gams et al. 2004). The Lecanoromycetes, 95 % of
which are lichenized fungi, are characterized by apothecioid
ascomata (rarely perithecioid) with an ascohymenial ontogeny
and a two-layered ascus wall with a rostrate dehiscence (Mi-
adlikowska et al. 2014, Gueidan et. al. 2015). Recent phylo-
genetic studies divide Lecanoromycetes into five subclasses:
Lecanoromycetidae, Ostropomycetidae, Umbilicariomycetidae,
Acarosporomycetidae and Candelariomycetidae (Hofstetter
et al. 2007, Miadlikowska et al. 2014). The subclass Ostro-
pomycetidae comprises the highest number of species with a
different lifestyle from the lichenized one (Baloch et al. 2010).
Several authors have proposed different hypotheses to explain
the evolution of the lichenicolous lifestyle. While Hawksworth
(1988) proposed that this lifestyle is just one more type of nutri-
tion within fungi, Lutzoni et al. (2001) put forward the idea that
the lichenicolous lifestyle originated from lichenized fungi and
that it is an intermediate stage towards other lifestyles, such
Phylogenetic placement within Lecanoromycetes
of lichenicolous fungi associated with Cladonia
and some other genera
R. Pino-Bodas1,2, M.P. Zhurbenko3, S. Stenroos1
1 Finnish Museum of Natural History, P.O.Box 7, FI-00014 University of
Helsinki, Finland; corresponding author e-mail: rpino@rjb.csic.es.
2 Real Jardín Botánico de Madrid (CSIC), Plaza Murillo 2, E-28014 Madrid,
Spain.
3 Komarov Botanical Institute, Russian Academy of Sciences, Professor
Popov 2, St. Petersburg, 197376, Russia.
Key words
cladoniicolous species
Pilocarpaceae
Protothelenellaceae
Scutula cladoniicola
Stictidaceae
Stictis cladoniae
Abstract Though most of the lichenicolous fungi belong to the Ascomycetes, their phylogenetic placement based
on molecular data is lacking for numerous species. In this study the phylogenetic placement of 19 species of
lichenicolous fungi was determined using four loci (LSU rDNA, SSU rDNA, ITS rDNA and mtSSU). The phylogenetic
analyses revealed that the studied lichenicolous fungi are widespread across the phylogeny of Lecanoromycetes.
One species is placed in Acarosporales, Sarcogyne sphaerospora; five species in Dactylosporaceae, Dactylo-
spora ahtii, D. deminuta, D. glaucoides, D. parasitica and Dactylospora sp.; four species belong to Lecanorales,
Lichenosticta alcicorniaria, Epicladonia simplex, E. stenospora and Scutula epiblastematica. The genus Epicladonia
is polyphyletic and the type E. sandstedei belongs to Leotiomycetes. Phaeopyxis punctum and Bachmanniomyces
uncialicola form a well supported clade in the Ostropomycetidae. Epigloea soleiformis is related to Arthrorhaphis
and Anzina. Four species are placed in Ostropales, Corticifraga peltigerae, Cryptodiscus epicladonia, C. galaninae
and C. cladoniicola comb. nov. (= Lettauia cladoniicola). Three new species are described, Dactylospora ahtii,
Cryptodiscus epicladonia and C. galaninae.
Article info Received: 31 October 2016; Accepted: 3 March 2017; Published: 28 June 2017.
92 Persoonia – Volume 39, 2017
Taxa Code Host species Voucher specimen ITS rDNA LSU rDNA SSU rDNA mtSSU
Bachmanniomyces uncialicola RP352 Cladonia stygia Finland, South Häme, R. Pino-Bodas s.n. (H) KY661637 – KY661702 –
RP123 Cladonia gracilis subsp. elongata USA, Alaska, Kodiak Island, S. & S. Talbot CHI017-56 (H) KY661617 – – –
Corticifraga peltigerae RP282 Peltigera elisabethae India, Jammu and Kashmir State, M.P. Zhurbenko 1353 (LE 260537) KY661634 KY661661 – KY661684
Cryptodiscus cladoniicola RP159 Cladonia furcata Czech Republic, Western Bohemia, J. Kocourková (H) KY661619 KY661652 KY661694 KY661674
(= Lettauia cladoniicola) RP160 Cladonia uncialis subsp. biuncialis Faroe Islands, Viðoy Island, J. Kocourková et al. s.n. (H) KY661620 KY661653 KY661695 KY661675
Cryptodiscus epicladonia RP208 Cladonia mitis USA, Alaska, Unimak, T. Ahti 70348a & S. Talbot (H) Holotype KY661628 – – KY661680
Cryptodiscus galaninae RP314 Cladonia rappii Canada, New Brunswick, T. Ahti 74421a & S. Clayden (H) KY661636 – KY661701 –
Dactylospora ahtii RP127 Cladonia gracilis USA, Alaska, Kodiak Island, S. & S. Talbot CHI017-63a (H) Holotype KY661618 – – –
RP182 Cladonia rangiferina USA, Alaska, Kodiak Island, S. & S. Talbot CHI17-37a (H) KY661622 – – KY661687
RP23 Cladonia uncialis subsp. biuncialis Iceland, Snæfellsnessýsla, F. Högnabba 1325a (H) KY661630 KY661659 – KY661686
Dactylospora deminuta RP235 Biatora vernalis Finland, Kuusamo Region, J. Pykälä 39390 (H) KY661629 – – KY661681
Dactylospora glaucomarioides RP275 Ochrolechia akagiensis Russia, Jewish Autonomous Region, M.P. Zhurbenko 13107 (LE 261065) KY661632 KY661660 – KY661683
Dactylospora parasitica RP422 Ochrolechia sp. Russia, Khabarovsk Territory, E.W. Tugi (LE 260868) KY661646 KY661666 – KY661690
RP423 Ochrolechia sp. Russia, Karachaevo-Cherkessia, M.P. Zhurbenko 12135 (LE 261336) – – – KY661691
RP424 Ochrolechia sp. Finland, Kuusamo Region, J. Pykälä 39145 (H) – KY661667 – KY661692
Dactylospora sp. RP391 Cladonia rangiferina Chile, Antártida chilena, W.R. Buck 60495a (H) – KY661664 – KY661689
Epicladonia sandstedei RP106 Cladonia coniocraea Finland, South Häme, Heinola, V. Haikonen 27543a (H) KY661614 KY661650 KY661693 KY661672
RP263 Cladonia sp. Russia, Republic of Adygeya, M.P. Zhurbenko 141 (LE 308482) KY661631 – – KY661682
Epicladonia simplex RP426 Cladonia botrytes Russia, Krasnoyarsk Territory, M.P. Zhurbenko 1050 (LE308685) KY661647 – – –
RP427 Cladonia botrytes Russia, Irkutsk Region, M.P. Zhurbenko 0563b (LE 309078) KY661649 – – –
RP428 Cladonia coccifera Russia, Tyumen’ Region, S.S. Kholod (LE 308573) KY661648 – – –
Epicladonia stenospora RP362 Cladonia humilis s. lat. Spain, Toledo, R. Pino-Bodas (H) – KY661663 KY661703 –
RP392 Cladonia pyxidata Turkey, Kars, M. Kocakoya 485a (H) – – KY661704 –
RP68 Cladonia rei Lithuania, Asveja Park, F. Högnabba 220911-15b (H) KY661640 KY661668 – –
RP119 Cladonia nana Portugal, Madeira, P. v.d. Boom 47938a (H) KY661616 KY661651 – KY661673
RP189 Cladonia chlorophaea USA, Alaska, Adak Island, S. & S. Talbot ADA 702a (H) KY661623 KY661654 – –
RP190 Cladonia pyxidata Turkey, Çankırı, M. Kocakaya 719b (H) KY661624 KY661655 KY661697 –
Epigloea soleiformis RP203 Cladonia subcervicornis Faroe Islands, Streymoy Island, J. Kocourková et al. s.n. (H) – – – KY661677
RP204 Cladonia subcervicornis Faroe Islands, Streymoy Island, J. Kocourková et al. s.n. (H) KY661625 KY661656 – KY661678
Lichenosticta alcicorniaria RP109 Cladonia pyxidata Russia, Russian Far East, J. Miadlikowska et al. KY661615 – – –
RP168 Cladonia arbuscula Finland, Uusimaa, R. Pino-Bodas s.n. (H) KY661621 – KY661696 KY661676
RP395 Cladonia tessellata Chile, Región de los Lagos, U. Schiefelbein (H) KY661638 KY661665 – –
Phaeopyxis punctum RP43 Cladonia ustulata New Zealand, S. Stenroos 6040a (H) KY661639 – – –
RP93 Cladonia coniocraea Finland, South Häme, V. Haikonen 29409 (H) KY661641 KY661669 – –
RP94 Cladonia coniocraea Finland, North Karelia, A. Launis 2212 (H) KY661642 – – –
RP95 Cladonia coniocraea Finland, North Karelia, A. Launis 2017 (H) KY661643 KY661670 – –
RP96 Cladonia coniocraea Finland, North Karelia, A. Launis 2213 (H) KY661644 KY661671 – –
RP97 Cladonia arbuscula Finland, Uusimaa, R. Pino-Bodas s.n. (H) KY661645 – KY661705 –
Protothelenella santessonii RP205 Cladonia subcervicornis Faroe Islands, Viðoy Island, J. Kocourková et al. s.n. (H) KY661626 KY661657 KY661698 KY661679
RP206 Cladonia subcervicornis Faroe Islands, Streymoy Island, J. Kocourková et al. s.n. (H) KY661627 KY661658 KY661699 –
Sarcogyne sphaerospora RP301 Candelariella sp. India, Jammu & Kashmir, Leh, M.P. Zhurbenko 1323 (LE 260996) KY661635 KY661662 KY661700 KY661685
Scutula epiblastematica RP276 Peltigera cf. malacea Russia, Sakha Republic, S.E. Vershinina (LE 261003) KY661633 – – KY661688
Table 1 List of specimens sequenced in this study, voucher information and the GenBank accession numbers.
93
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
as saprophytism or parasitism. If the latter hypothesis was
true, we would expect a greater number of lichenicolous fungi
to belong to Lecanoromycetes. Moreover, it is worth pointing
out that according to some studies the lichenicolous lifestyle
is more flexible than was thought (Wedin et al. 2004). Many
optionally lichenicolous species are known, such as several
species of the genus Chroodiscus (Lücking & Grube 2002) or
Diploschistes muscorum, which in the initial stages of develop-
ment parasitizes Cladonia species and subsequently forms an
independent lichenized thallus (Friedl 1987).
The present work mainly focuses on the lichenicolous fungi that
live on Cladonia (Lecanorales, Ascomycota), a sub-cosmopoli-
tan genus with 470 species (Ahti pers. comm.) characterized by
a dimorphic thallus formed by a primary crustose or squamulose
thallus and a fruticulose secondary thallus (Ahti 2000). Cur-
rently, 128 species of obligately lichenicolous fungi are known
to live on Cladonia, which is one of the lichen host genera along
with Lecanora, Peltigera and Pseudocyphellaria on which most
species of lichenicolous fungi have been reported (Hawksworth
& Miadlikowska 1997, Lawrey & Diederich 2016, Zhurbenko &
Pino-Bodas 2017). Some authors proposed that certain gen-
era, such as Peltigera or Pseudocyphellaria are suitable hosts
for the development of lichenicolous fungi because they have
large thalli and live in damp habitats (Etayo & Diederich 1996,
Etayo & Sancho 2008). This explanation can also be applied
to the genus Cladonia that can form wide mats and cover the
soil in areas where humidity is rather high. The cladoniicolous
species of Lecanoromycetes occur in the genera Dactylospora,
Diploschistes, Phaeopyxis, Protothelenella, Scutula and Stictis
(Lumbsch & Huhndorf 2011, Suija et al. 2015), the phylogenetic
positions of which has been confirmed by molecular data only
for the optionally lichenicolous Diploschistes muscorum and for
Phaeopyxis punctum (Suija et al. 2015). The aim of this study
was to determine the phylogenetic placement of 19 species of
lichenicolous fungi, most of which live on species of the genus
Cladonia, using four loci.
MATERIALS AND METHODS
Material studied and taxon sampling
Specimens of Cladonia species from the herbaria H and LE
plus new collections (also deposited in H or LE) were screened
in order to find lichenicolous fungi. In addition, lichenicolous
fungi on other lichen genera were selected to complete the
sampling. The morphology and anatomy of the specimens were
examined and photographed using dissecting microscopes
Stemi 2000-CS and Leica DFC490, and compound micro-
scopes Axio Imager A1 (equipped with Nomarski differential
interference contrast optics) and Leica DM2500. Microscopic
examination was done in water, 10 % KOH (K), Meltzer, Lu-
gol’s iodine, directly (I) or after a KOH pre-treatment (K/I), or
phloxine. The length, breadth and length/breadth ratio (l/b) of
asci and ascospores are given (where n > 10) as: (minimum–)
{X–SD} –{X +SD}(– maximum), where X is the arithmetic mean
and SD the corresponding standard deviation, followed by the
number of measurements.
For the molecular study 74 fresh specimens were selected.
Unfortunately, for many specimens the amplifications were
not successful, and DNA sequences were obtained only from
43 specimens, representing 19 species (Table 1). We tried to
select at least two specimens per species studied, but only one
specimen could be sequenced for some of the species (Table 1),
owing to the difficulties of finding additional fresh material or to
the amplification failure of additional specimens.
The DNA sequences were first included in the dataset of Schoch
et al. (2009), allowing us to verify that all species in the pre-
sent study belong to Lecanoromycetes (data not shown). For
the phylogenetic analyses the sampling was completed with
sequences downloaded from GenBank (Appendix 1), based
on the results of Miadlikowska et al. (2014) plus sequences of
lichenicolous fungi belonging to the Lecanoromycetes (Lawrey
& Diederich 2016). The clades containing the species studied
were sampled more intensively. Leotia lubrica was selected
as outgroup. Several species were placed in the family Sticti-
daceae, and separate phylogenetic analyses were run for this
family based on the phylogenies of Baloch et al. (2009, 2013).
DNA extraction, PCR and sequencing
Lichen thalli were cleaned by Milli-Q SP Ultra-Pure-Water, then
1–10 lichenicolous ascomata or conidiomata were removed
using an insect needle size 00 (Imperial Karlsbad) and cleaned
from the remaining lichen using a sterilized razor blade. Geno-
mic DNA was extracted using E.Z.N.A. Forensic DNA Isolation
Kit (Omega BioTek). DNA was eluted in the final step in 100
µl of elution buffer provided by the manufacturer. Four loci
were selected to infer the phylogeny: ITS rDNA, LSU rDNA,
SSU rDNA and mtSSU. In addition, RPB1 and RPB2 were
also tested, with different combinations of primers (gRPB1-Af/
gRPB1-CR, RPB2-607F/RPB2-1554R, RPB2-5F/ RPB2-7R,
fRPB27cF/fRPB211aR) but the amplifications were not suc-
cessful. The PCRs were carried out using ReadytoGoPCR
Beads (GE Healthcare Life Sciences, UK), with 25 µl of final
volume, 1 µl of each primer at 10 µM concentration and 3 µl of
DNA. The primers used were: ITS1F/ITS4 (White et al. 1990,
Gardes & Bruns 1993) for ITS rDNA; mrSSU1/mrSSU3R (Zoller
et al. 1999) for mtSSU; LROR/ LR5 or LR6 (Vilgalys & Hester
1990, Vilgalys & Sun 1994) for LSU rDNA; and NS1/NS22 or
NS24 (White et al. 1990, Gargas & Taylor 1992) for SSU rDNA.
Amplifications were performed using an Eppendorf Master
cycler ep Gradient S thermal cycler with the following programs:
95 °C 5 min; 5 cycles of 30 s at 95 °C, 30 s at 58 °C, 60 s at
72 °C; 30 cycles of 30 s at 95 °C, 30 s at 56 °C, 60 s at 72 °C;
7 min at 72 °C for ITS rDNA; 95 °C 5 min; 30 cycles of 30 s at
95 °C, 30 s at 55 °C, 60 s at 72 °C; 10 min at 72 °C for LSU
rDNA; 95 °C 5 min; 40 cyles of 30 s at 95 °C, 40 s at 52 °C,
60 s at 72 °C; 10 min at 72 °C for SSU rDNA; 95 °C 5 min; 35
cycles of 30 s at 95 °C, 60 s at 50 °C, 60 s at 72 °C; 7 min at
72 °C for mtSSU. PCR products were cleaned with GFX PCR
DNA and Gel Band Purification kit (GE Healthcare), E.Z.N.A.
UltraSep Gel Extraction Kit (Omega BioTek), or Illustra TM
ExoProStar TM 1step (GE Healthcare). Sequencing was
performed at Macrogen Europe service (www.macrogen.com).
Phylogenetic analyses
The sequences were assembled in Sequencher 4.1.4 program
(Gene Codes Corporation, Inc, Ann Arbor, Michigan, USA).
BLAST searches (Altschul et al. 1997, www.ncbi.nlm.nih.gov/
BLAST) were done for each sequence in order to dismiss
contaminations and to check which taxa are most similar to
our sequences.
The sequences were aligned using MAFFT (Katoh & Standley
2013) with different algorithms depending on the input, then
the alignments were improved manually in BIOEDIT 7.0 (Hall
1999). Introns and ambiguous regions were removed from the
alignments with Gblock 0.91b (Castresana 2000) using the
less stringent option.
Each dataset was analyzed with maximum likelihood (ML)
analysis in RAxML 7.0.3 (Stamatakis et al. 2005), using the
GTRGAMMA model and with 500 replicates of fast bootstrap in
order to check conflicts among the datasets, following Hillis &
Bull (1993) criteria. No incongruence was found and the data-
sets were combined. The optimal substitution model for each
locus (Table 2) was selected with jModeltest (Posada 2008)
94 Persoonia – Volume 39, 2017
using the Akaike Information Criterion (AIC). The combined
dataset was analyzed with ML and Bayesian inference (BI).
The ML analysis was run in RAxML considering each locus
as different partition with the GTRGAMMA model and 1 000
replicates of fast bootstrap to assess the node support. The
Bayesian analysis was run in MrBayes 3.2.6 (Ronquist et al.
2012) in CIPRES Science Gateway v. 3.1 (Miller et al. 2010).
The posterior probabilities were approximated by sampling trees
using Markov Chain Monte Carlo (MCMC). Two simultaneous
runs with 90 000 000 generations each, starting with a random
tree and employing 6 simultaneous chains, were executed.
Every 2 000th tree was saved into a file. The convergence was
assessed in Tracer v. 1.5 (Rambaut & Drummond 2009) plot-
ting the likelihood versus generation number and the average
standard deviation of split frequencies (≤ 0.01). The first 50 %
trees were discarded as burn-in and the consensus tree was
calculated with the remaining 22 500 trees.
Additionally, a phylogeny of the family Stictidaceae, based on
LSU rDNA, mtSSU and ITS rDNA, was constructed to study
more accurately the relationship of Lettauia and Cryptodiscus
species. Trapeliopsis flexuosa and Xylographa parallela were
used as outgroup. For every dataset an ML analysis was run
according to the options above. The datasets were congruent
and they were combined. The combined dataset was analyzed
with ML, considering each locus as different partition with the
GTRGAMMA model and 1 000 replicates of fast bootstrap to as-
sess the node support. The optimal substitution model for each
locus was selected with jModeltest, these models are listed in
Table 2. The Bayesian analysis was run with two simultaneous
runs of 10 000 000 generations each, starting with a random
tree and employing 4 simultaneous chains. The convergence
was assessed with the same method as in the previous analy-
sis. The initial 50 % trees were discarded as burnin and the
consensus tree was calculated.
Topological hypothesis tests
The phylogenetic analyses revealed placements or taxa cir
cumscriptions incongruent with the current classifications,
whereby alternative phylogenetic topologies were tested: a) the
monophyly of the genus Epicladonia; b) the genus Lettauia
belongs to the family Fuscidiaceae. First, the optimal ML trees
were estimated in RAxML using the GTRGAMMA model and
considering each locus as a different partition. Shimodaira-
Hasegawa test (SH, Shimodaira & Hasegawa 1999) and
expected likelihood weight (ELW, Strimmer & Rambaut 2002)
were conducted in TREEPUZZLE 5.2 (Schmidt et al. 2002),
using the GTR+I+G model with fourcategory approximation to
the gamma distribution for substitution rate among sites and
using 1 000 RELL bootstrap replicates.
RESULTS
In this study, 92 new sequences were generated (36 of ITS
rDNA, 22 of LSU rDNA, 21 of mtSSU and 13 of SSU rDNA).
Members of the genera Bachmanniomyces, Corticifraga, Epi-
cladonia, Epigloea, Lettauia and Lichenosticta were sequenced
for the first time in this study.
BLAST searches revealed a similarity between the sequences
generated here and the ones deposited in GenBank. The
results are listed in Appendix 2. The most similar sequences
corresponded to Lecanoromycetes sequences or, in some
cases, to sequences coming from nonidentified environmen-
tal fungi. BLAST searches revealed that the sequences most
similar to Epicladonia sandstedei corresponded to sequences
of the Leotiomycetes. The mtSSU sequence of Dactylospora
deminuta showed an 85 % similarity with one sequence of
the Chaetothyriales. The BLAST searches did not generate
similarity with sequences of the genus Cladonia or with any
other host genus. Therefore we can maintain that none of the
sequences included in the analyses corresponds to the host.
Table 2 summarizes the data for single loci datasets. The con-
catenated dataset included 285 sequences and 3 264 charac-
ters. The ML analysis yielded a tree with LnL = 106201.732,
while the Bayesian analyses yielded a consensus tree with
LnL = 102147.35 (arithmetic mean). The ML tree and the
Bayesian consensus had a similar topology. The Bayesian
consensus tree is shown in Fig. 1. The general topology agreed
with the recently published phylogenies of the Lecanoromycetes
(Miadlikowska et al. 2006, 2014), showing the same main clad-
es (although some of them were not supported). According to
our phylogenetic analyses one lichenicolous species belonged
to the Acarosporales, Sarcogyne sphaerospora (Fig. 1); it was
phylogenetically related to Polysporina subfuscescens with high
support. Four species were included in the order Ostropales,
Corticifraga peltigerae, Cryptodiscus epicladonia, Cryptodiscus
galaninae and Lettauia cladoniicola, (Fig. 1). Corticifraga pelti-
gerae is closely related to Actinoplaca strigulacea in the family
Graphidaceae, subfamily Gomphilloideae (Fig. 1). Lettauia
cladoniicola and the two new species of Cryptodiscus were
placed in the Stictidaceae (Fig. 1). Four species were placed
in the order Lecanorales, Epicladonia simplex, E. stenospora,
Lichenosticta alcicorniaria and Scutula epiblastematica (Fig. 1).
The three specimens of Lichenosticta alcicorniaria formed a
wellsupported clade. This clade turned out to be phylogeneti-
cally related to Gypsoplaca macrophylla, but the relationship
lacked support in all the analyses. The genus Epicladonia was
polyphyletic, the type species E. sandstedei was monophyletic
(two specimens studied) but it fell outside the class Lecanoro-
mycetes. The other two species, E. stenospora and E. simplex
formed a well-supported clade inside the family Pilocarpaceae,
possibly related to the genus Micarea (low statistical support).
N bp NV NP CI/RI Model -Lnl
Lecanoromycetes
LSU rDNA 226 1270 837 592 0.1838 /0.5391 GTR+I+G 39343.844280
SSU rDNA 184 1019 386 305 0.2672 /0.5578 GTR+I+G 15289.101485
ITS rDNA 156 379 297 248 0.1632/ 0.5057 HKY+I+G 16141.468421
mtSSU 227 596 484 414 0.1563/0.5991 GTR+I+G 31424.666287
Stictidaceae
LSU rDNA 27 870 288 200 0.6167/0.7235 GTR+I+G 4373.206559
ITS rDNA 20 477 262 195 0.5711/0.5942 GTR+I+G 3916.270036
mtSSU 28 702 325 267 0.5366/0.7100 GTR+I+G 5234.960566
Table 2 Features of each dataset analyzed, including number of sequences aligned (N), number of positions in each aligment (bp), number of variable
positions (NV), number of parsimony informative positions (NP), consistence index (CI), retention index (RI), model of evolution selected with jmodeltest and
likelihood from ML analysis.
95
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
Arthrorhaphis citrinella
Dactylospora deminuta RP235
Dactylospora parasitica RP423
Cudoniella clavus
Maronea chilensis
Stictis urceolatum
Cryptodiscus galaninae RP314
Cryptodiscus epicladonia RP208
Sphaeropezia ochrolechiae
Absconditella sphagnorum
Pleopsidium gobiense
Protothelenella santessonii RP206
Stictis radiata
Carestiella socia
Dactylospora mangrovei
Ropalospora chlorantha
Petractis nodispora
Phaeopyxis punctum RP94
Thelenella antarctica
Fuscidea austera
Thrombium epigaeum
Chlorociboria aeruginosa
Protothelenella corrosa
Pleopsidium chlorophanum
Acarospora laqueata
Epigloea soleiformis RP204
Phaeopyxis punctum 3
Epicladonia sandstedi RP263
Cryptodiscus pallidus
Phaeopyxis punctum RP96
Phaeopyxis punctum RP93
Sarcogyne plicata
Dactylospora ahtii RP23
Botryotinia fuckeliana
Protothelenella sphinctrinoidella
Dactylospora haliotrepha
Sphaeropezia mycoblasti
Dactylospora sp. RP391
Acarosporina microspora
Sarcogyne hypophaea
Polysporina arenacea
Polysporina subfuscescens 2
Lettauia cladoniicola RP160
Bachmanniomyces uncialicola RP352
Dactylospora vrijmoediae
Ingvariella bispora
Fuscidea cyathoides
Protothelenella santessonii RP205
Sclerococcum sphaerale 1
Epigloea soleiformis RP203
Sarcogyne sphaerospora RP301
Cryptodiscus foveolaris
Phaeopyxis punctum RP95
Mollisia cinerea
Epicladonia sandstedei RP106
Anzina carneonivea
Bachmanniomyces uncialicola RP123
Cryptodiscus gloeocapsa
Dermea acerina
Sclerococcum sphaerale 2
Sarcogyne clavus
Sarcogyne algoviae
Phaeopyxis punctum RP43
Dactylospora glaucomarioides RP275
Schizoxylon albescens
Sarcogyne regularis
Phaeopyxis punctum RP97
Phaeopyxis punctum 2
Polysporina subfuscescens 1
Phaeopyxis punctum 1
Leotia lubrica
Xyloschistes platytropa
Umbilicariaceae
(6)
Pertusariales
(6)
Ostropales
Trapeliales
(6)
Arctomiales (6)
Baeomycetales (3)
Hymeneliales (2)
Stictidaceae
Dactylospora parasitica RP422
Dactylospora parasitica RP424
Dactylospora ahtii RP127
Dactylospora ahtii RP182
Dactylosporaceae
Acarosporaceae
Lettauia cladoniicola RP159
Protothelenellaceae
-/0.97
91/-
-/0.97
79/-
-/0.97
Umbilicariales
Acarosporales
Ophioparmaceae (3)
Leotiomycetes
Geoglossomycetes (2)
Lichinomycetes (2)
Candelariales (2)
-/0.98
100/-
-/0.97
-/0.97
-/0.97
Epigloeaceae
Fig. 1 This is the 50 %majorityrule consensus tree of Bayesian analysis of Lecanoromycetes based on nLSU, nSSU, mtSSU and ITS rDNA. Branches sup-
ported with posterior probability ≥ 0.95 and bootstrap ≥ 70 % are indicated in bold. Grey rectangles show the groups where lichenicolous fungi studied were
placed. Lichenicolous fungi are marked with a black circle. The black triangles indicate lichenicolous lichens. The squares mark the facultative lichenicolous
species. The bold names indicate the newly sequence specimens (extraction codes are indicated). Classification according to Miadlikowska et al. (2014).
96 Persoonia – Volume 39, 2017
Gyalecta jenensis
Phlyctis argena
Porpidia speirea
Corticifraga peltigerae RP282
Tetramelas pulverulentus 2
Fissurina insidiosa
Lecidoma demissum
Porpidia albocaerulescens
Solenopsora candicans
Lecidea fuscoatra
Tetramelas phaeophysciae 2
Fellhanera bouteillei
Tetramelas pulverulentus 1
Tetramelas phaeophysciae 1
Porina lectissima
Gyalidea hyalinescens
Diploschistes muscorum
Micarea denigrata
Actinoplaca strigulacea
Puttea margaritella
Cecidonia xenophana
Lecidea auriculata
Epicladonia simplex RP427
Lecidea laboriosa
Byssoloma subdiscordans
Diploschistes euganeus
Calenia monospora
Micarea denigrata
Lecidea silacea
Graphis scripta
Cecidonia umbonella
Psilolechia leprosa
Micarea alabastrites
Diploschistes cinereocaesius
Coenogonium luteum
Myriotrema olivaceum
Platythecium grammitis
Immersaria usbekica
Tephromela atra
Sagiolechia protuberans
Petractis clausa
Scoliciosporum intrusum
Calopadia foliicola
Bellemerea alpina
Fissurina sp.
Mycoblastus sanguinarius
0.95
0.95
Peltigerales
(16)
Caliciales
(6)
Teloschistales
(22)
Rhizocarpales (3)
Ostropales
Gomphilloideae
Epicladonia simplex RP426
Epicladonia simplex RP428
Epicladonia stenospora RP189
Epicladonia stenospora RP119
Epicladonia stenospora RP68
Epicladonia stenospora RP190
Epicladonia stenospora RP362
Epicladonia stenospora RP392
Lecanorales
81/-
83/-
100/-
-/0.97
75/-
Lecideales
Pilocarpaceae
Fig. 1 (cont.)
97
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
Fig. 1 (cont.)
Both species of Epicladonia (E. simplex and E. stenospora)
were monophyletic. Scutula epiblastemica was placed in the
Ramalinaceae, it was related to S. miliaris and S. tuberculosa.
Other species were included in different families with uncer-
tain phylogenetic placement in the Lecanoromycetes (Dac-
tylosporaceae, Epigloeaceae and Protothenellaceae). Five
species were placed in the family Dactylosporaceae (Fig. 1),
Dactylospora ahtii, D. deminuta, D. glaucomarioides, D. para-
sitica (the generic type) and Dactylospora sp. Three specimens
of D. parasitica formed a well-supported clade together with
Sclerococcum sphaerale. The three specimens of the new
species Dactylospora ahtii were monophyletic. Dactylospora
glaucomarioides grouped with Dactylospora sp. Protothelenella
santessonii was monophyletic and formed a well-supported
clade with the other species of Protothelenella (Fig. 1). Epigloea
soleiformis was placed in the Ostropomycetidae and is related
to the genera Arthrorhaphis and Anzina (Fig. 1). Phaeopyxis
punctum and Bachmanniomyces uncialicola were included in
the Ostropomycetidae but their relationships within this subclass
were not resolved.
The combined dataset of the Stictidaceae contained 2 049
characters, the ML analysis yielded a tree with a likelihood value
of -LnL = 13788.449, while the arithmetic mean likelihood of
Bayesian analysis was LnL = 14304.77. The topology of both
trees was the same and so only the Bayesian 50 % consensus
majority tree is shown (Fig. 2). The genus Lettauia and two new-
ly described species clustered in the genus Cryptodiscus, with
high support (100 % of bootstrap/1.00 of posterior probability).
The Cryptodiscus clade is closely related to a clade formed by
Ingvariella bispora and Xyloschistes platytropa. Acarosporina
microspora, Carestiella sociata, Ostropa barbata, Schizoxylon
albescens, Stictis confusa and S. populorum formed another
wellsupported clade. The genus Stictis was polyphyletic. The
genera Absconditella, Geisleria and Sphaeropezia turned out
to be closely related. The SH and ELW tests rejected both
alternative hypothesis tested (Table 3).
0.09
Crocynia pyxinoides
Scutula miliaris
Lichenosticta alcicorniaria RP395
Carbonea supersparsa
Punctelia rudecta
Miriquidica garovaglii 2
Spaerophorus fragilis
Lopezaria versicolor
Dactylina arctica
Alectoria ochroleuca
Carbonea vitellinaria
Lecanora hybocarpa
Flavocetraria nivalis
Biatora subduplex
Nesolechia oxyspora 2
Bacidina arnoldiana
Carbonicola anthracophila
Psora decipiens
Pycnothelia papillaria
Cetraria islandica
Lecanora conizaeoides
Hypogymnia physodes
Lecania cyrtella
Sphaerophorus globosus
Lichenosticta alcicorniaria RP109
Carbonea vorticosa
Pseudephebe pubescens
Protoblastenia rupestris
Xanthoparmelia conspersa
Phacopsis vulpina
Cladonia stipitata
Parmelina tiliacea
Platismatia glauca
Adelolecia pilati
Lecidella elaeochroma
Miriquidica garovaglii 1
Lecanora contractula
Cladonia caroliniana
Protoblastenia calva
Lepraria lobificans
Protoparmelia atriseda
Mycobilimbia tetramera
Parmotrema tinctorum
Ramalina complanata
Lichenosticta alcicorniaria RP168
Hypotrachyna degelii
Ramboldia elabens
Scutula epiblastematica RP276
Protoparmelia phaeonesos
Melanelia fuliginosa
Biatora alaskana
Usnea antarctica
Lecanora strobilina
Gypsoplaca macrophylla 2
Ramboldia gowardiana
Gypsoplaca macrophylla 1
Imshaugia aleurites
Raesenenia huuskonenii
Protoparmelia cupreobadia
Evernia prunastri
Vulpicida pinastri
Lecanora achariana
Flavoparmelia caperata
Nesolechia oxyspora 1
Ramalina farinacea
Scutula tuberculosa
Ramboldia insidiosa
Bryoria trichodes
Bacidia schweinitzii
Rhizoplaca melanophthalma
Lecanorales
Ramalinaceae
99/-
86/-
100/-
-/0.97 -/0.97
-/0.96
-/0.97
-/0.98
-/0.99
100/-
99/-
92/-
98 Persoonia – Volume 39, 2017
Geisleria sychnogonoides
Sphaeropezia capreae 1
Sphaeropezia mycoblasti
Cryptodiscus pallidus
1
Stictis confusum
Cryptodiscus pallidus 2
Xyloschistes platytropa
Schizoxylon albescens
Stictis radiata
Sphaeropezia lyckselensis
Lettauia cladoniicola RP159
Cryptodiscus foveolaris
Absconditella sphagnorum 1
Cryptodiscus tabularum 1
Stictis populorum
Carestiella sociata
Sphaeropezia capreae 2
Cryptodiscus tabularum 2
Xylographa parallela
Cryptodiscus gloeocapsa 1
Cryptodiscus epicladonia RP208
Cryptodiscus gloeocapsa 2
Stictis urceolatum
Cryptodiscus pini
Absconditella sphagnorum 2
Trapeliopsis flexuosa
Cryptodiscus galaninae RP314
Ingvariella bispora
Ostropa barbara
Acarosporina microspora
Lettauia cladoniicola RP160
97/1
99/1
100/1
100/1
100/1
100/1
100/1
100/1
100/1
100/1
100/1
100/1
76/1
100/1
100/1
100/1
-/1
92/1
90
/1
-/0.98
100/1
77/-
93/0.99
100/0.99
-/0.99
-/0.99
0.05
Fig. 2 Phylogeny of the family Stictidaceae. This is the 50 %majorityrule
consensus tree of a Bayesian anlysis based on nLSU, mtSSU and ITS
rDNA. Posterior probability ≥ 0.95 and bootstrap ≥ 70 % are indicated on
the branches. The bold names indicate the newly sequence specimens
(extraction codes are indicated).
TAXONOMY
Cryptodiscus cladoniicola (D. Hawksw. & R. Sant.) Pino-
Bodas, Zhurb. & S. Stenroos, comb. nov. — MycoBank
MB820201; Fig. 3
Basionym. Lettauia cladoniicola D. Hawksw. & R. Sant., Biblioth. Lichenol.
38: 138. 1990.
Type. Germany, Baden, Schwarzwald, Feldberg-Gipfels, Nordseite,
elev. 1400 m, on Cladonia amaurocraea (podetia), 14 July 1912, G. Lettau,
holotype B 7700.
Ascomata apothecia, soon sessile, more or less round in sur-
face view, slightly constricted at the base, 180– 300(– 430) µm
diam, disc initially plane, pale yellow/orange yellow, becoming
convex (up to hemispherical) and light to moderate orange
under aging, epruinose, margin initially slightly raised, white,
20–40(–60) µm wide, becoming lacerated, reduced or even
excluded under aging; dispersed or occasionally aggregated to
contiguous. Proper exciple composed of round or tangentially
elongated cells c. 2.5–6 × 2–3 µm with walls 0.5 –1 µm thick,
without embedded crystals; lateral exciple hyaline except for
the light orange yellow outermost part, 25–40 µm thick; lower
exciple (hypothecium) hyaline, 15–40 µm thick. Periphysoids
absent. Epihymenium light orange yellow, c. 5 µm tall. Hyme-
nium hyaline, 30–50 µm tall, I + fleetingly blue then immediately
yellow green (mainly due to yellow colouration of ascal plasma)
with some remnants of blue colouration, K/ I+ blue or partly
red due to colouration of ascal walls. Subhymenium hyaline,
c. 10 µm tall. Paraphyses filiform, often di or occasionally tri-
chotomically branched, mainly above, 1.2–1.7(–3.0) µm diam,
frequently septate, often somewhat constricted at the septa
and strangulated, particularly near the apices, which are occa-
sionally slightly swollen. Asci narrowly clavate to subcylindrical,
with short foot, (43–)44 –48(–50) × 6.5–9 (–10) µm (n = 16, in
water, I or K/I), tholus up to 5 µm tall, I–, K/I–, apical structures
not observed, wall/periascal gel I+ fleetingly blue, K/I+ blue or
partly red, 8-spored. Ascospores hyaline, cylindrical to slightly
fusiform, the apices rounded or occasionally acute, (13.5–)
16.8–22.8(–26.0) × (2.0–)2.3 –2.9(–3.5) µm, l/b = (4.9–)
6.3–8.9(–12.2) (n = 54, in water, I or K), (2–)3(–4)-septate,
not constricted at the septa, wall smooth, without a gelatinous
sheath, with conspicuous guttules, arranged in the ascus in a
bundle, diagonally or overlappingly 2–4 seriate. Anamorph not
found.
Distribution & Hosts — The species is known from Austria,
the British Isles, Canada, the Czech Republic, Denmark,
Finland, Germany, Norway, Russia, Slovenia, Sweden and
the USA, growing on podetia of Cladonia amaurocraea, C. ar-
buscula, C. furcata, C. gracilis, C. mitis, C. portentosa, C. rangi-
ferina, C. stellaris, C. stygia and C. uncialis (Hawksworth &
Santesson 1990, Alstrup 1993, Coppins 1998, Diederich 2003,
Santesson et al. 2004, Kocourková & Van den Boom 2005,
Hafellner 2008, present paper). Cladonia uncialis is a new host
species. Pathogenicity not observed.
Specimens examined. CzeCh republiC, Western Bohemia, distr. Karlovy
Vary, Bečov nad Teplou, 1 km E of the town, Psí skála hill, on Cladonia furcata
(podetia), 2 Aug. 2014, J. Kocourková, H. – Denmark, Faroe Islands, Viðoy
Hypothesis -Lnl SH ELW
Monophyly of Epicladonia 111117.04 0.0010* 0.0000*
Lettauia belongs to Fuscidiaceae 111260.55 0.0000* 0.0000*
* indicate significant results.
Table 3 Results of topological tests Shimodaira-Hasegawa (SH) and likeli-
hood weight test (ELW).
99
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
Fig. 3 Cryptodiscus cladoniicola. a. Appearance of apothecia: a1–a3 from LE 308679, a4 from LE 308798, a5 f from LE 308695; b. apothecial section in water
from LE 308798; c. exciple and hymenium in cross section in I from LE 308798; d. asci in water from LE 308696; e. asci in K/ I from LE 308695; f– g. paraphyses
in K from LE 308679; h. paraphyses in K from LE 308695; i. ascospores in K, i1 from LE 308679, i2 from LE 308695. — Scale bars: a = 200 µm; b = 20 µm;
c– i = 10 µm.
Fig. 4 Cryptodiscus epicladonia. a. Appearance of apothecia from the holotype; b. lateral exciple in cross section in K from LE 338773a; c. asci in water from
the holotype; d. apothecial section in water from LE 308773a; e. ascus in I from the holotype; f. paraphyses in phloxine from LE 308498; g. ascospore in water
from LE 308498. — Scale bars: a = 500 μm; b– c, e–g = 10 μm; d = 50 μm.
100 Persoonia – Volume 39, 2017
Island, Viðareiðy, Mýrnafjall Mt, 3 km SE of the town, Bergshálsur plateau
on north end of the mountain crest, on C. uncialis (podetia), 13 Aug. 2013,
J. Kocourková, W.J. Halda & I. Sommerová, H. – russia, Krasnoyarsk Ter-
ritory, Putorana Plateau, Kapchuk Lake, on C. arbuscula (podetia), 18 Aug.
1983, M.P. Zhurbenko 83236, LE 308897; Krasnoyarsk Territory, Western
Sayan Mts, Ergaki Nature Park, Olen’ya River, on C. arbuscula (podetia), 11
July 2010, M.P. Zhurbenko 1041, LE 308679; ibid., on C. mitis (podetia), 11
July 2010, M.P. Zhurbenko 1053, LE 308684; Republic of Sakha (Yakutia),
Indigirka River, Silyapskii Range, on C. rangiferina (bases of podetia), 24
June 1976, I.I. Makarova, LE 308798; Primorye Territory, SikhoteAlin’ Range,
Mt Glukhomanka, on C. uncialis (podetia), 21 Aug. 2003, K.S. Podlubnaya,
LE 308695.
Notes — There are some discrepancies with the detailed
species description in Hawksworth & Santesson (1990) who
reported more or less plane apothecia up to 250 µm diam,
an I+ blue hymenium up to 65 µm tall, sometimes anastomo-
sed paraphyses and (1–)3-septate ascospores, measuring
19–25(–31) × 2.5 – 3 µm. The species was formerly reported in
Russia from Bol’shezemel’skaya tundra in Nenets Autonomous
Area (LE 210357, Zhurbenko 2008), the Northern Ural Mts in
Komi Republic (LE 308521, Zhurbenko 2004) and Putorana
Plateau in Krasnoyarsk Territory (LE 207133, Zhurbenko 2000).
We confirm the identification of LE 308521, while LE 210357
belongs to Cryptodiscus galaninae; the identification of LE
207133 is uncertain due to scanty material.
Cryptodiscus epicladonia Zhurb. & Pino-Bodas, sp. nov. —
MycoBank MB820198; Fig. 4
Etymology. Referring to its occurrence on Cladonia.
Type. usa, Alaska, Aleutian Islands, Unimak Is., False Pass, 3 km SW
of airstrip, N54.837° E163.417°, elev. 160 m, on Cladonia mitis (podetia),
25 Aug. 2011, T. Ahti & S. Talbot 70348a, holotype H.
Diagnosis — Lichenicolous fungus. Differs from Stictis clado-
niae mainly in the light orange yellow with white pruinose rim
vs brownish black and epruinose ascomata, the hyaline to very
pale orange yellow vs mainly medium to dark brown proper
exciple, the I–, K/I– vs I+ red, K/I+ blue hymenium, the longer
asci, mainly 73–93 × 7– 9 µm and the longer, (5–)7–11-septate
ascospores, mainly 50–73 × 1.5– 2 µm.
Ascomata apothecioid, more or less superficial, initially al-
most closed, later widely urceolate, roundish, hemispherical,
broader or narrower at the base, 100–500 µm diam, 50 –160
µm tall, laterally light orange-yellow, above usually with a
white, coarsely granulose, sometimes outwardly extending
crystalline rim 20–80 µm wide; disc concolorous with lateral
parts, sometimes slightly more intensively coloured, rounded to
elongated in surface view, 50–100 µm lengthways; scattered to
aggregated, sometimes adjacent. Proper exciple composed of
thick-walled, rounded or somewhat elongated cells c. 2–6 µm
lengthways; lateral exciple 40–100 µm thick, hyaline, outwardly
usually covered by 10–40 µm thick layer of colourless crystals
2–12 µm across; lower exciple (hypothecium) 20– 30 µm thick,
hyaline to very pale orange yellow at the base. Periphysoids
absent. Epihymenium indistinct. Hymenium hyaline, 70–100
µm tall, I–, K/I–. Subhymenium hyaline, 10 –30 µm thick, com-
posed of thin-walled more or less isodiametric cells c. 2–4 µm
diam, hardly distinct from lower exciple. Paraphyses filiform,
septate, 0.8–1.5 µm diam, apices usually spathulate or capi-
tate, occasionally shortly forked, 1.5–3.0 µm diam, sometimes
protruding above the hymenium. Asci subcylindrical to elongate
clavate, with short foot, apex rounded, tholus 1–3(– 9) µm thick,
sometimes with a narrow apical beak to 2 µm tall, (71–)73–
93(–97) × (6 –)7–9 µm (n = 13, in water, phloxine, I or K/I),
I–, K/I–, 8-spored. Ascospores hyaline, filiform to cylindrical,
slightly tapering towards the apices, (37.0–)50.0–72.5(–87.0)
× (1.3–)1.5–1.9(–2.2) µm, l/b = (22–)28 –46(– 55) (n = 81, in
Characters C. cladoniicola C. epicladonia C. galaninae Stictis cladoniae
Apothecia plane to convex urceolate urceolate urceolate
disc pale yellow, orange-yellow or orange disc light orange-yellow disc pale to yellow or yellowish white disc blackish
margin white margin orange-yellow margin subhyaline or yellowish white margin brownish black
epruinose white pruinose rim epruinose epruinose
Lateral exciple (thickness) 25 ‒40 µm 40‒100 µm 15 ‒40 µm 20‒ 70 µm
Epihymenium light orange yellow indistinct indistinct indistinct
Hymenium 30‒ 50 µm tall 70‒100 µm tall 30‒45 µm tall 50‒70 µm tall
I+ fleetingly blue then immediately yellow I‒, K/I‒ I+ blue then quickly orange to red I+ red, K/I+ blue
green, K/I+ blue or partly K/I+ blue with occasional reddish stripes
Asci (mainly) 44‒ 48 × 6.5‒9 µm 73‒ 93 × 7‒9 µm 28‒38 × 5‒ 6.5 µm 57‒71 × 8‒10 µm
Ascospores cylindrical to slightly fusiform filiform to cylindrical fusiform, slightly clavate or bacilliform filiform to cylindrical,
7‒23 × 2.5 ‒3 µm 50‒73 × 1.5 ‒2 µm 9.5 ‒12.5 × 1.5‒2 µm c. 40 ‒60 × 1.5‒2 µm
(2‒) 3(‒ 4)septate (5‒)7‒11septate (0‒)1(‒ 2)septate at least 4‒ 5septate
Table 4 Main morphological differences among the cladoniicolous species of the genus Cryptodiscus and Stictis cladoniae (based on original data).
101
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
water, phloxine, I or K/I), (5 –)7–11-septate (septa sometimes
indistinct), not constricted at the septa, smooth-walled, lacking
a gelatinous sheath, with many small, hardly conspicuous gut-
tules, arranged in the ascus in a bundle. Anamorph not found.
Distribution & Hosts — The species is known from tundra
(mainly) and taiga biomes of Asia and North America, growing
on podetia and rarely basal squamules of Cladonia amauro-
craea, C. arbuscula, C. mitis and C. uncialis. Pathogenicity not
observed.
Additional specimens examined. CanaDa, Newfoundland & Labrador,
Labrador Straits, L’ Anse Amour, on Cladonia arbuscula (podetia), 10 Sept.
2015, T. Ahti 75728a & J.M. McCarthy, H. – russia, Krasnoyarsk Territory,
Taimyr Peninsula, Osipovka, on C. arbuscula (podetia), 18 July 1990, M.P.
Zhurbenko 901105, LE 308498; same peninsula, Levinson-Lessing Lake,
on C. arbuscula (moribund bases of podetia), 27 Aug. 1995, M.P. Zhurbenko
95598, LE 308913; Chukotka Autonomous Area, Provideniya, on C. uncialis
(bases of podetia), 3 July 1969, Safronov, LE 308789; Chukotka Autonomous
Area, lower Bol’shoi Anyui River, on C. amaurocraea (bases of podetia), 11
July 1951, V.N. Andreev, LE 308795; Chukotka Autonomous Area, head-
waters of Utesiki River, on C. amaurocraea (podetia), 21 July 1948, M.N.
Avramchik, LE 308773a.
Notes — With respect to the other cladoniicolous fungi,
Cryptodiscus epicladonia morphologically resembles C. clado-
niicola, C. galaninae and Stictis cladoniae, which are compared
in Table 4.
Cryptodiscus galaninae Zhurb. & Pino-Bodas, sp. nov. —
MycoBank MB820199; Fig. 5
Etymology. The species is named after the Russian lichenologist Irina A.
Galanina, who collected the type.
Type. russia, Magadan Region, Ol’skii District, km 82 of road Magadan-
Talon, near Magtur field station, N59°45'27" W149°39'56", elev. 26 m, on
Cladonia sp. (moribund podetia), 7 Aug. 2013, I.A. Galanina, holotype LE
308693.
Diagnosis — Lichenicolous fungus. Differs from Cryptodiscus
foveolaris in the I+ red and shorter hymenium 30–45 µm vs
Fig. 5 Cryptodiscus galaninae. a. Appearance of apothecia from the holotype; b. apothecia, b1 from the holotype, b2– b5 from LE 380730; c. apothecial section
in water from LE 308741; d. apothecial section in I from the holotype; e. short-celled hyphae on the inner side of lateral exciple in K from LE 309139; f. asci
in K/I from LE 309139; g. ascus in K from the holotype; h. asci in K/I from the holotype; i. ascospores in K from the holotype. — Scale bars: a –b = 200 µm;
c = 50 µm; d– i = 10 µm.
102 Persoonia – Volume 39, 2017
50–80 µm tall, the shorter asci 27– 42 µm vs 50 –65 µm long,
the longer and narrower ascospores 7–14.5 × 1.5–2 µm vs
6–9 × 2.5–3 µm and in the lichenicolous life habit.
Ascomata apothecioid, initially immersed in the host thallus
then erumpent and eventually superficial, cupulate, round to
ellipsoid in surface view, sometimes constricted at the base,
widely urceolate, epruinose, up to 330 µm diam, up to 150 µm
tall; margin subhyaline or yellow-white, up to 40 µm thick;
disc deeply concave, pale to moderate yellow or yellow-white,
translucent, glossy; usually aggregated to contiguous. Proper
exciple composed of isodiametric or tangentially elongated
cells 2–6 × 1– 4 µm with walls 0.5–2 µm thick, hyaline, not dif-
ferentiated into layers, without embedded crystals, 15–40 µm
thick laterally, 5–10 µm thick below the hymenium. Periphysoids
absent, but short-celled hyphae reminiscent of those mentioned
in Baloch et al. (2009: 60) have been observed on the inner side
of lateral exciple (Fig. 5d). Epihymenium indistinct. Hymenium
hyaline, 30–45 µm tall, I+ blue then quickly orange to red, K/I+
blue with occasional reddish stripes. Subhymenium hyaline,
c. 5 µm tall. Paraphyses filiform, septate, mainly 1.2–1.5 µm
diam, up to 2.5 µm diam at the base and up to 2 µm diam at
the apices, which are sometimes slightly clavate, rarely with
short branchlets or forked in the upper part. Asci subcylindrical
to elongate clavate, with short foot, apex rounded, tholus up to
2.5 µm thick, apical structures not observed, (27–)28 –38(–42)
× (4.5–)5 –6.5(–7) µm (n = 20, in water, I, K or K/I), I –, periascal
gel K/ I+ blue, 8-spored. Ascospores hyaline, slightly fusiform
or slightly clavate (tapering down), occasionally almost bacilli-
form, straight, (7.1–)9.6 –12.4(–14.5) × (1.3–)1.6 – 2.0(– 2.2)
µm, l/b = (4.2 –)5.0–7.4(–9.7) (n = 140, in water, I, K or K/I),
(0–)1(–2)-septate, not constricted at the septa, with thin and
smooth wall, lacking a gelatinous sheath, sometimes with con-
spicuous guttules, diagonally or overlappingly 2–4-seriate in
the ascus. Anamorph not found.
Distribution & Hosts — The species is known from tundra
and taiga biomes of Europe, Asia and North America, growing
on aged or moribund podetia or rarely basal squamules of
Cladonia gracilis, C. rangiferina, C. rappii s.lat., C. umbricola
and Cladonia sp. Pathogenicity not observed.
Additional specimens examined. CanaDa, British Columbia, Columbia Mts,
Beaver River, on Cladonia umbricola (basal squamules), 17 July 2002, M.P.
Zhurbenko 02100c, LE 308741; British Columbia, Wells Gray Provincial Park,
Mt Raft, on C. rangiferina (podetia), 3 Aug. 2002, M.P. Zhurbenko 02309, LE
308730; New Brunswick, Charlotte Co., 1.5 km NNW of Chance Harbour
along power line corridor W of Route 790, on C. rappii s.lat. (moribund
podetia), 6 Sept. 2014, T. Ahti 74421a & S.R. Clayden, H. – russia, Nenets
Autonomous Area, Bol’shezemel’skaya tundra, Khar’yaga oilfield, on C. rangi-
ferina (podetia), 25 July 2007, M.P. Zhurbenko 0735, LE 210357 (formerly
erroneously identified and published as Lettauia cladoniicola (Zhurbenko
2008)). – usa, Alaska, Kotzebue, on C. gracilis (moribund podetia), 19 Aug.
2000, M.P. Zhurbenko 00239, LE 309139.
Notes — Cryptodiscus galaninae is quite distinct from the
other species of the genus with 1-septate ascospores, viz.
C. foveolaris and C. pini (Baloch et al. 2009). Both of these
species are saprotrophs on wood, the former one can be dis-
tinguished by its I– and taller hymenium 50–80 µm tall, longer
asci 50–65 × 4 –5 µm and shorter and wider ascospores 6 –9 ×
2.5–3 µm; the latter one differs in its larger ascomata 0.3 –0.6
mm diam with dark reddish brown outer layer of the exciple,
I– and taller hymenium 60 –80 µm tall and larger asci 40–60 ×
6–7 µm. The other known species of Cryptodiscus also growing
on Cladonia are C. cladoniicola and C. epicladonia described
here in detail. The differences among these species are present-
ed in Table 4. The other lichenicolous fungi with urceolate apothe-
cia and hyaline ascospores growing on Cladonia are Biazrovia
stereocaulicola, Spirographa fusisporella and Stictis cladoniae.
Biazrovia stereocaulicola can easily be distinguished from Cryp-
todiscus galaninae by its vinaceous, cinnamon or orange-
brown apothecia and ellipsoid, 3-septate, larger ascospores
measuring (12–)15– 20(–28) × (4 –)4.5–5.5(–6.5) µm (Zhur-
benko & Etayo 2013). Spirographa fusisporella is distinct by
its 16–32-spored asci and helicoid, longer ascospores 22 –31
× 1–2.5 µm (Diederich 2004). The differences from Stictis
cladoniae can be found in Table 4.
Dactylospora ahtii Zhurb. & Pino-Bodas, sp. nov. — Myco-
Bank MB820200; Fig. 6
Etymology. The species is named after the Finnish lichenologist, our
friend Prof. Teuvo Ahti.
Type. usa, Alaska, Kodiak Archipelago, Chirikof Island, N55.77095°
W155.63464°, elev. 174 m, on Cladonia gracilis subsp. vulnerata (podetia),
19 July 2013, S. & S. Talbot CHI017-67a, holotype H.
Diagnosis — Lichenicolous fungus. Differs from Dactylo spora
aeruginosa mainly in the stipitate ascomata, the shorter hy-
menium, 40–60 µm tall, somewhat smaller ascospores, (7.6 –)
10.4–13.0(–16.3) × (3.0–)3.5–4.3(–5.5) µm vs (9–)11–14.5
(–16) × (3–)3.5–5.5(–7) µm and in the disparate hosts.
Ascomata apothecia, more or less scattered, composed of a
disc usually sitting on a distinct stipe (in LE 264407 stipe poorly
developed); disc shiny, dark brown to almost black when dry,
medium brown and somewhat translucent when wet, epruinose,
round, plane to somewhat concave, occasionally urceolate
in senescent overmature apothecia with disintegrated hyme-
nium, (80–)130– 250(–600) µm diam (n = 103), surrounded
by a usually slightly elevated, often darker (particularly when
wet) distinct margin, in side view forming a sharply delimited
marginal flange 15–40 µm thick protruting from the stipe for
20–40 µm; stipe usually somewhat tapering towards the base,
typically 80–230 µm wide, 40 –100 µm tall, pale to medium
brown, much paler than the disc or occasionally concolo rous.
Proper exciple 15–30(–70) µm thick laterally, up to 150 µm tall
basally, where it forms a stipe; consists of a cupulate, medium
red-brown or orange-brown inner layer and of a subhyaline
or pale red-brown to orange-brown outer layer with a darker
red-brown to orange-brown outermost edge c. 5 µm thick;
the outer layer composed of comparatively large isodiametric
to tangentially elongated cells c. 5–11 µm lengthways, with
walls 1–2 µm thick; the upper lateral part of the exciple usu-
ally contains deep purple to dark violet, K+ dark green to
blue-green blotches (not observed in LE 264407). Epihyme-
nium unevenly pale to medium red-brown to orange-brown,
pigmentation amorphous, 5(–10) µm tall, sometimes rather
indistinct. Hymenium hyaline to pale red or orange-brown,
40–60 µm tall, I+ blue above, red below or I+ blue throughout
(in LE 308774), K/I+ blue with red patches. Apothecial section
K– (except for the blotches) or becomes less red. Paraphyses
septate, somewhat constricted at the septa, particularly above,
occasionally with ramifications above, 1.5 –2 µm diam, api-
cal cells usually medium red or orange-brown, more or less
capitate, 3–4(–5.5) µm diam, sometimes not pigmented and
only slightly enlarged. Asci elongate clavate, c. 40–55 × 9 –12
µm, 8-spored, with I+ blue, K/I+ blue external gelatinous cap,
8-spored. Ascospores hyaline or rarely light brown, homopolar
to somewhat heteropolar, ellipsoid to slightly obovate (with a
wider upper cell), occasionally oblong, straight or occasionally
slightly curved, (7.6–)10.4–13.0 (–16.3) × (3.0–)3.5 –4.3(–5.5)
µm, l/ b = (1.8–)2.6–3.4(– 4.3) (n = 302, in water, K, I or K/I),
(0–)1-septate, not or occasionally slightly constricted at the
septum, guttulate, wall c. 0.5 µm thick, smooth, without internal
thickenings, non-halonate, arranged irregularly 2–3-seriate in
the ascus. Anamorph not found.
Distribution & Hosts — The species is known from polar
desert, tundra (mainly) and taiga biomes of Europe, Asia and
North America, growing on podetia of Cladonia arbuscula,
103
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
C. gracilis subsp. vulnerata, C. mitis, C. portentosa subsp.
pacifica, C. rangiferina (most finds) and C. uncialis. Often
grows on aged parts of host podetia, visible damage to the
host not observed.
Additional specimens examined. GreenlanD, Frederikshåbs Isblink,
on Cladonia rangiferina (podetia), 7 July 2009, E.S. Hansen, Lichenes
Groenlandici Exsiccati 1092a, H; Siorapaluk, on C. rangiferina (podetia), 25
July 2009, E.S. Hansen, H. – iCelanD, Snæfellsnessýsla, Fróðárheiði pass,
between Mt Miðfell and Mt Knarrarfjall, on C. rangiferina and C. uncialis
(podetia), 22 July 2009, F. Högnabba 1325c, H. – norway, Svalbard, Alde-
gondabreen glacier, on C. rangiferina (podetia), 16 July 2003, M.P. Zhur-
benko 03211, LE 264407. – russia, Murmansk Region, Khibiny Mts, Mt Kukis-
vumchorr, on C. rangiferina (base of podetia), 9 Aug. 1997, M.P. Zhurbenko
971, LE 207408 (formerly erroneously reported as Scutula epicladonia in
Zhurbenko 2001); Krasnoyarsk Territory, Severnaya Zemlya Archipelago,
Bol’shevik Is., Mt Bol’shaya, on C. rangiferina (podetia), 27 Aug. 1998, N.V.
Matveeva, LE 308885; Krasnoyarsk Territory, Taimyr Peninsula, mouth of
Pyasina River, on C. rangiferina (base of podetia), 6 Aug. 1993, V.B. Kuvaev
2184, LE 207407 (formerly erroneously reported as Scutula epicladonia in
Zhurbenko & Santesson 1996); same peninsula, Levinson-Lessinga Lake,
on C. rangiferina (moribund bases of podetia), 28 July 1995, M.P. Zhurbenko
95592, LE 308880; same peninsula, Kotui River, Kayak, on C. rangiferina
(podetia), 24 July 1996, I.Yu. Kirtsideli, LE 308937; Republic of Sakha
(Yakutia), Indigirka River, Ust’-Nera, on C. rangiferina (podetia), 11 July
1992, M.P. Zhurbenko 92568, LE 308922; Chukotka Autonomous Area,
Innepinkuliveem River, on C. mitis (podetia), 10 Aug. 1951, Ababkov, LE
308796; Chukotka Autonomous Area, Lorino, on C. arbuscula (bases of
podetia), 16 Aug. 1972, I.I. Makarova, LE 308781. – USA, Alaska, Seward
Peninsula, 7 km ESE of Nome, on C. rangiferina (moribund podetia),
1 Sept. 2001, M.P. Zhurbenko 0142c, LE 308589c, M.P. Zhurbenko 0171,
LE 308516; Mause Creek, on C. rangiferina (podetia), 22 July 2000, D.A.
Walker, LE 309135; Kotzebue, on C. rangiferina (podetia), 19 Aug. 2000,
M.P. Zhurbenko 00232, LE 309138; Kobuk Valley Wilderness, Waring Mts,
on C. arbuscula (podetia), 31 July 2000, M.P. Zhurbenko 00139, LE 309137;
Kodiak Archipelago, Chirikof Is., on C. rangiferina (podetia), 19 July 2013,
S. & S. Talbot CHI017-65a, H; Aleutian Islands, Carlisle Is., on C. gracilis
subsp. vulnerata (podetia), 28 July 2013, S. & S. Talbot CAR001-23b, H;
same islands, northwest corner of Amalia Is., on C. gracilis subsp. vulnerata
(podetia), 2 Aug. 2013, S. & S. Talbot AML305a, H; same islands, Adak Is.,
northern side of Finger Bay, on C. rangiferina (podetia), 26 Aug. 2013, S. &
S. Talbot ADA717a, H; Wosnesenski Is., Port Moller, on C. portentosa subsp.
pacifica (podetia), 31 June 2009, S. Talbot WOS019-19a, H.
Notes — Compared to the Dactylospora species with 1-sep-
tate ascospores produced in 8-spored asci D. ahtii is most
similar to Dactylospora sp. (presented below), D. aeruginosa
and D. protothallina. The Dactylospora sp. differs from D. ahtii
in having only occasionally stipitate ascomata with a much
shorter stipe, a completely dark reddish orange or brown
upper part of the exciple, more intensively red tinge of epi-
hymenium and proper exciple and K+ aeruginose blotches
sometimes located in the hypothecium. Further, Dactylospora
sp. differs in its ascospores, which are constantly pale to me-
dium pigmented, somewhat larger, (8.9–)10.9 –14.9(–18.3)
× (3.4–)4.5–5.9(–7.6) µm, exceptionally also 2-septate and
sometimes distinctly constricted at the septum. Dactylospora
aeruginosa differs from the new species in having non-stipitate
apothecia, a much thicker hymenium mainly 70–120 µm thick,
violet-blue, K+ aeruginose blotches occurring not only in the
lateral exciple, but also in the epihymenium and hymenium,
a light brown hypothecium and somewhat larger ascospores,
(9–)11–14.5(–16) × (3 –)3.5–5.5(–7) µm, with a perispore up
to 2 µm thick (Ihlen et al. 2004). This species have been re-
ported from the coastal forests of Norway, Alaska and from the
Arctic, growing on thalli of various epiphytic crustose lichens
from the genera Biatora, Japewia, Lopadium and Micarea
or directly on wood and bark of Picea and Juniperus and on
terricolous crustose lichens Lecidea epiphaea (Zhurbenko &
Von Brackel 2013) and Biatora subduplex (as ‘cf.’; Zhurbenko
2009). Dactylospora protothallina differs from D. ahtii in the
absence of K+ aeruginose blotches, a brown epihymenium, a
somewhat taller hymenium of 65–80 µm and somewhat wider,
brown ascospores (9–)10–15 × 4.5 –7.5 µm (Hafellner 1979,
Nimis 1993, Alstrup & Ahti 2007, Spribille et al. 2010). So far,
D. prothallina has been reported from the lichen species of
Fuscopannaria, Massalongia, Parmeliella, Protopannaria and
from adjacent biofilms. The other Dactylospora species reported
Fig. 6 Dactylospora ahtii. a. Appearance of apothecia, a1–a2 from LE207408, a3 –a4 from LE 207407, a5 from the holotype; b. apothecial section in water
from LE 207408; c. dark purple excipular blotches in water from LE 207407; d. hymenium in K from the holotype; e. asci in K/ I from the holotype; f. ascospores
in K from the holotype. — Scale bars: a = 200 µm; b = 20 µm; c– f = 10 µm.
104 Persoonia – Volume 39, 2017
on Cladonia are D. cladoniicola, so far known only from the
holotype on Cladonia macrophyllodes collected in Svalbard
(Alstrup & Olech 1993) and D. deminuta, a widely distributed
species recorded from many unrelated host genera. Both of
them have brown mature ascospores. In addition, D. cladoni-
icola has much larger ascospores measuring 33–37 × 12 –14
µm and D. deminuta has (3–)5 –7(–8)-transseptate ascospores.
Another similar species is Scutula cladoniicola, which differs
from Dactylospora ahtii in the following characters:
1. apothecial stipe usually absent or, if present, shorter than
40 µm and concolorous with the disc;
2. apothecial disc blackish and not translucent when wet;
3. lateral exciple medium brown throughout, with hyaline
outermost edge;
4. epihymenium indistinct;
5. hymenium hyaline to olive grey below;
6. violet, K+ aeruginose blotches occur also in the hymenium;
7. apices of paraphyses usually medium reddish orange-
brown, more or less capitate, 3–4(–5.5) µm diam;
8. amyloid external gelatinous cap of the asci not observed;
9. ascospores hyaline, usually homopolar, larger, 13.0–16.4
× 5.5–6.7, (0 –)1(– 3)-septate, with granulate wall 0.5–1
µm thick.
Dactylospora sp. — Fig. 7
Apothecia blackish, glossy, 0.2– 0.6 mm diam, sessile, without a
stipe or with a short paler stipe up to 40 µm tall (in LE 308774),
disc plane to convex, margin thin, prominent, concolorous with
the disc, not translucent when wet. Epihymenium medium
red-brown, to 10 µm tall. Paraphyses with somewhat swol-
len apices 3–4 µm diam. Hymenium more or less colourless,
40–60 µm tall. Proper exciple red-orange or orange-brown,
dark and 25–50 µm thick laterally and pale (but with darker
marginal rim), 40–50 µm thick below the hymenium, where it
is composed of much larger, mainly isodiametric cells up to
16 µm across with relatively thin wall. Lower exciple (hypothe-
cium) medium to dark red-brown, up to 100 µm tall, with dark,
indistinctly coloured, K+ aeruginose blotches or without them
(in LE 308809). Apothecial section becomes less reddish in
K. Asci 8-spored. Ascospores pale yellow-gray-olive-brown
to medium brown, slightly obovate (with wider upper cell) to
occasionally ellipsoid, (0–)1(–2)-septate (only exceptionally
aseptate or 2-septate), sometimes distinctly constricted at the
septum, (8.9–)10.9–14.9(–18.3) × (3.4–)4.5 –5.9(–7.6) µm,
l/b = (1.6 –)2.1–2.9(–3.9) (n = 256, in water, I or K), smooth,
non-halonate.
Distribution & Hosts — The species is known from tundra
and taiga biomes of Asia and from the subantarctic part of
South America. Mainly found on moribund parts of Cladonia
amaurocraea, C. cariosa, C. rangiferina and C. symphycarpa,
but also occur on adjacent biofilms and plant remnants and
thus probably somewhat saprobic.
Specimens examined. Chile, Antártida Chilena, Comuna Cabo de Hornos,
Alberto de Agostini National Park, Hoste Is., on Cladonia rangiferina (pode-
tia), 16 Jan. 2013, W.R. Buck 60495a, H (specimens sequenced). – russia,
Krasnoyarsk Territory, Eastern Sayan Mts, Kryzhina Range, Belyi Kitat River,
on C. symphycarpa (moribund basal squamules) and biofilms over terrico
lous mosses, 14 July 2009, M.P. Zhurbenko 0956, LE 308658; Republic of
Sakha (Yakutia), Olenek Region, Siibikte River basin, on C. cariosa (basal
squamules) and occasionally on adjacent plant remnants, 11 Aug. 1957, A.N.
Lukicheva, LE 308809; Chukotka Autonomous Area, Pekul’nei Range, on
C. amaurocraea (moribund base of podetia), 4 July 1950, M.N. Avramchik,
LE 308774.
Notes — The examined material resembles Dactylospora
ahtii, D. aeruginosa and D. protothallina. Dactylospora aerugi-
nosa can be distinguished by its much taller hymenium (up to
120 µm), a light brown hypothecium and halonate ascospores
(Ihlen et al. 2004). Dactylospora protothallina differs in the
absence of K+ aeruginose blotches and a brown epihymenium
(Hafellner 1979). The differences with D. ahtii have been dis-
cussed above under the latter species. The studied specimens
might represent a new species of Dactylospora, but it is not
formally described, pending the discovery of additional material.
Scutula cladoniicola Alstrup & D. Hawksw., Meddel. Gronland,
Biosci. 31: 65. 1990 — Fig. 8
Type. GreenlanD, Near Ivigtut, N61°14', elev. 0– 50 m, on the ground in
dwarf shrub heath, on Cladonia stricta (podetia), 9 July 1946, M.S. Chris-
tiansen 5504, holotype herb. Christiansen, C (?), isotype IMI 331024! The
type host is apparently Cladonia trassii, not C. stricta, which was misused
in 1946.
Ascomata apothecial, sessile, black throughout, not translucent
when wet, epruinose, glossy, rounded, strongly constricted at
the base to short stipitate, 150–800 µm diam, up to 450 µm tall,
disc plane, somewhat convex or concave, margin slightly raised
or flush with the disc. Lateral exciple 40–60(–100) µm thick,
moderate brown, K+ brown-orange, with hyaline outermost lay-
er c. 5 µm thick, in cross section composed of radially elongated
cells c. 5.5–17 × 4– 9 µm, with walls 1–3 µm thick. Lower exciple
(hypothecium) up to 350 µm tall, merging with lateral exciple,
moderate brown, K+ brownish orange, in cross section com-
posed of rounded cells with walls 1.5–4 µm thick. Epihymenium
indistinct. Hymenium (40–)50 –70 µm tall, hyaline throughout
or olive grey below, with scattered orange yellow crystalline
granules on the surface, I+ blue, K/ I+ blue with occasional red
patches. K+ blue-green blotches are scattered in lateral exciple
(mainly), lower exciple and hymenium. Paraphyses 1.8– 2.9 µm
diam, apices reddish orange-brown, slightly clavate, 2.5–3.2
µm diam, septate, sometimes slightly constricted at the septa
(particularly in K), occasionally branched and anastomosed.
Asci narrowly clavate, c. 40–65 × 8 –11 µm, staining of tholus
with I and K/I not observed, but periascal gel I and K/I+ blue,
8-spored. Ascospores hyaline, usually homopolar, ellipsoid,
occasionally oblong or rarely obovate, (10.0–)13.0–16.4 (–19.0)
× (4.5–)5.5– 6.7(–7.5) µm, l/b = (1.7–)2.0 –2.8(– 3.6) (n = 152,
in water, I, K or K/I), (0 –)1(–2 or exceptionally – 3)-septate, not
constricted at the septum, wall 0.5–1 µm thick, granulate, lack-
ing a gelatinous sheath, overlappingly uniseriate to irregularly
biseriate in the ascus.
Distribution & Hosts — The species was reported from
the Arctic Canada, Greenland, Iceland and Turkey (Alstrup &
Hawksworth 1990, Hansen & Alstrup 1995, Von Brackel 2010,
Zhurbenko 2013, Kocakaya et al. 2016), growing on Cladonia
monomorpha, C. pyxidata, C. rangiferina and C. stricta.
Fig. 7 Dactylospora sp. (LE 308658) a. Apothecial section in water; b. vari-
ation of ascospores in water. — Scale bars: a = 20 µm; b = 10 µm.
105
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
Notes — There are some discrepancies between the ex-
amined isotype of the species and its protologue (Alstrup &
Hawksworth 1990), where anastomoses of the paraphyses
and violet blotches in the proper exciple and hymenium were
not mentioned, the epihymenium was reported being greyish
brown and interspersed with greenish granules, the apical cells
of paraphyses bearing a brown gelatinous coat, the asci with
I+ structures in the tholus (Alstrup & Hawksworth 1990: f. 38C)
and the ascospores 1(–2)-septate, (12.5–)13 –15(–16) × 5 –6.5
µm. Morphologically, Scutula cladoniicola recalls Dactylospora
ahtii, their distinguishing characters being summarized under
the latter species.
Stictis cladoniae (Rehm) Sacc., Syll. Fung. (Abellini) 8: 692.
1889 — Fig. 9
Type. austria, Tyrol, Piztal valley, near the Taschach glacier, elev. c. 2000
m, on Cladonia gracilis s.l. (podetia), Aug. 1875, H. Rehm, holotype S! The
type host is apparently Cladonia macroceras, since C. gracilis should not
occur in Tyrol.
Ascomata apothecioid, initially immersed and almost or possibly
completely closed, later superficial and deeply urceolate, up to
530 µm diam and up to 190 µm tall, brown-black, epruinose, with
blackish disc, scattered. Proper exciple cupulate, 20–70 µm
thick laterally, 15– 25 µm thick basally, uniformly medium to dark
brown except for the pale brown to hyaline internal lateral parts,
composed of thick-walled, tangentially more or less elongated
cells. Periphysoids absent. Epihymenium indistinct. Hymenium
hyaline, 50–70 µm tall, I+ red, K/I+ blue. Subhymenium hyaline,
composed of thin-walled isodiametric cells, up to 10 µm tall.
Paraphyses filiform, unbranched, septate, 1.5–1.7 µm diam,
apices somewhat enlarged, to 2.4 µm diam. Asci subcylindri-
cal, with short foot, endoascus thickened at the apex to 2.5
µm, apical beak not observed, (55–)57–71(–78) × (7–)8–10
µm (n = 17, in K/I), periascal gel I+ red, K/ I+ blue, staining of
apical structures in K/I not observed, 8-spored. Ascospores
hyaline, filiform/cylindrical, slightly tapering towards the apices,
c. 40–60 × 1.5 – 2 µm (n = 13, in K/I), septation was obscure,
but 4–5-septate spores were observed, smooth-walled, without
a perispore, guttulate, arranged in the ascus in a bundle.
Notes — So far the species was known only from the type
collection in the Austrian Alps (Rehm 1882) and from Lappland
in Sweden (Santesson et al. 2004), growing on Cladonia gra-
cilis s.lat. We revised its holotype, as the former descriptions
of the species (Saccardo 1889, Rehm 1912, Sherwood 1977)
essentially recapitulate the protologue (Rehm 1882), where
its ascospores were described as being aseptate and much
shorter, c. 36 × 1.5– 2 µm, and asci shorter, c. 48– 50 × 8 µm.
DISCUSSION
Prior studies have proved that the lichenicolous lifestyle arose
multiple times along biological evolution (Arnold et al. 2009,
Diederich et al. 2013, Suija et al. 2015). Within Lecanoromycet-
es, Divakar et al. (2015) showed that the lichenicolous lifestyle
originated at least three times in the family Parmeliaceae. The
results presented here confirm manifold independent origins
of the lichenicolous lifestyle in the class Lecanoromycetes.
The richest order as for lichenicolous fungi is the Lecanorales,
followed by the Ostropales (Fig. 1). In addition, the latter com-
prises the greatest number of species with a different lifestyle
from the lichenized one within Lecanoromycetes (Schoch et al.
2009, Baloch et al. 2010).
The family Dactylosporaceae was introduced by Bellemere &
Hafellner (1982) to fit the genus Dactylospora. The species of
this genus are characterized by a type of asci with an I– tholus
covered by an I+ blue external gelatinous cap (Hafellner 1979,
Bellemere & Hafellner 1982). Molecular studies have shown
different phylogenetic positions for this family, while in the phy-
logenies published by Schoch et al. (2009) and Diederich et al.
(2013) it was placed in the Eurotiomycetes. Miadlikowska et al.
Fig. 8 Scutula cladoniicola (isotype). a. Apothecia on host surface; b. appearance of apothecia; c. apothecial section in K; d. ascomatal section in K/I;
e. paraphyses in K; f. ascus in K/I; g. ascospores in K; h. ascospores in K/I. — Scale bars: a–b = 0.5 mm; c = 50 µm; d = 20 µm; e –g = 10 µm.
106 Persoonia – Volume 39, 2017
Sarcogyne sphaerospora was placed in the family Acaro-
sporaceae, related with Polysporina subfuscescens (Fig. 1).
This result agrees with its current classification based on mor-
phology (Hafellner 1995). The genera Sarcogyne and Polyspo-
rina differ in the presence of a carbonized epihymenium in the
latter (Vězda 1978). Lendemer et al. (2009) pointed out that
this character could be insufficient to keep these genera apart.
The recent phylogenetic analyses of the family Acarosporaceae
(Reeb et al. 2004, Westberg et al. 2015) indicate that both
genera are polyphyletic and that a carbonized epihymenium
is not restricted to a unique phylogenetic lineage (Westberg
et al. 2015). In turn it has been shown that Polysporina sub-
fuscescens is a polyphyletic species (Westberg et al. 2015).
On the basis of morphology, it has been considered that
S. sphaerospora could be related to Acarospora stapfiana and
A. succedens (Lendemer et al. 2009). These two species share
with S. sphaerospora the presence of spherical ascospores
with a perispore. This relationship is highly probable since
other species of Sarcogyne and Acarospora have been shown
to be closely related (Westberg et al. 2015). The family Acaro-
sporaceae needs an exhaustive taxonomical study in order to
delimit the genera and the species.
In the phylogenetic analysis presented by Suija et al. (2015) a
common cladoniicolous fungus, Phaeopyxis punctum (type spe-
cies of the genus) was placed in the Lecanoromycetes, subclass
Ostropomycetidae, but its relationships within this subclass was
not resolved. Our results, based on the sequences of six new
specimens, confirm the placement of P. punctum in the Lecano-
romycetes but do not solve either the relationship of the species
within Ostropomycetidae. Our phylogenetic analyses showed
that P. punctum along with the coelomycete Bachmanniomyces
uncialicola (also confined to species of Cladonia) form a well-
supported clade on the base of the Ostropomycetidae (Fig. 1).
Phaeopyxis punctum frequently grows on both podetia and
basal squamules of Cladonia and usually does not induce galls,
while Bachmanniomyces uncialicola mostly grows on podetia,
(2014) suggested that the family belongs to Lecanoromycetes,
a more consistent result with the morphological characters
of the genus, and confirmed in our analyses. These authors
recommend using more than six loci to obtain a well-founded re-
sult about the phylogenetic position of the family. Since we have
not sequenced additional genes we do not discuss the phylo-
genetic position of the family and limit ourselves to describe the
relationships between the species. Schoch et al. (2009) found
that the genus Dactylospora was polyphyletic, and that the de-
termination of which of the two clades represented Dactylospora
s.str. was still pending. In the present work three specimens
of the generic type species, D. parasitica, were included and
we confirmed that this species belongs to the clade formed by
D. haliotrepha and D. mangrovei in the phylogeny of Schoch
et al. (2009). Dactylospora parasitica, formed a well-supported
monophyletic group with the sporodochial hyphomycete Scle-
rococcum sphaerale (Hawksworth 1975, Diederich et al. 2013).
Both species mainly grow on species of the genus Pertusaria.
Excluding this relationship, the species with the same lichen
host genera were not phylogenetically related. Dactylospora
ahtii and Dactylospora sp., both growing on the genus Cla-
donia, are not closely related. Dactylospora sp. is related to
D. glaucomarioides growing on Ochrolechia. Dactylospora glau-
comarioides morphologically resembles D. parasitica (Hafellner
1979), while Dactylospora sp., is more similar to D. aerugi-
nosa (species not studied here). Dactylospora ahtii resembles
D. aeruginosa and D. protothallina (see above). Dactylospora
deminuta represents an early-diverging lineage in the genus,
apparently with a very long branch. This could be due to the
fact that we only achieved sequencing two loci (ITS rDNA and
mtSSU). The ancestor of the family could have a lichenicolous
lifestyle and the switch to saprobic lifestyle have occurred in
the lineage formed by D. mangrovei, D. haliotrepha and D. vrij-
moediae. But this hypothesis must be proved with more loci
and more species, since in our phylogenetic analyses most of
the relationships among species are not supported.
Fig. 9 Stictis cladoniae (holotype). a. Appearance of apothecia; b. apothecial section in K; c. hymenium and basal exciple in K; d. asci with spores in K/I;
e. asci with spores and lateral exciple in K/I. — Scale bars: a = 0.5 mm; b = 20 µm; c –e = 10 µm.
107
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
only rarely on basal squamules and almost always induces
galls (Zhurbenko & Pino-Bodas 2017). However, gall formation
has also been reported for Phaeopyxis punctum (Grummann
1960, Rambold & Triebel 1990, Zhurbenko & PinoBodas 2017),
and occasionally both species grow together (Motiejūnaitė
et al. 2011, our own specimens on Cladonia stygia, Finland,
R. Pino-Bodas s.n., H). The two binomials may refer to the
same species, as indicated by our phylogenetic analyses, and
B. uncialicola may be an anamorph of Phaeopyxis punctum.
So far the phylogenetic placement of the genus Epigloea was
uncertain in the Ascomycota. Davis (1987) created the family
Epigloeaceae, exclusively containing the genus Epigloea. The
features peculiar to Epigloea are gelatinous perithecioid asco-
mata, nonfissitunicate, 8 to multispored asci with an I+ wall
and colourless septate ascospores sometimes with terminal
apiculae (Döbbeler 1984, Davis 1987, Pérez-Ortega & Bar-
reno 2006). Originally the genus was considered as lichenized
(Zukal 1890), but later Döbbeler (1984) showed it to be a
highly specialized parasite of algae. One species, Epigloea
urosperma, is exclusively lichenicolous, and two other species,
E. bactrospora and E. soleiformis, occasionally grow on lichens
(Döbbeler 1994, Zhurbenko 2010, Czarnota & Hernik 2013).
No author has found morphological characters that permit to
place this genus in some of the groups of the Ascomycota.
Our phylogenetic analyses show that E. soleiformis belongs to
the subclass Ostropomycetidae, close to Anzina carneonivea
and Arthrorhaphis citrinella. The placement of Epigloea in
the Ostropomycetidae is not particularly surprising, because
this class comprises species with different types of ascomata
(Grube et al. 2004, Schmitt et al. 2005, 2009). Nevertheless,
no morphological character suggested beforehand that this
genus could be related to the genera Anzina or Arthrorhaphis.
However, the confirmation of the phylogenetic position of the
genus Epigloea will require the inclusion of the type species,
E. bactrospora, in a phylogenetic study.
The family Protothelenellaceae was first placed in the Ostropo-
mycetidae by Schmitt et al. (2005). We have sequenced for the
first time one of the three known lichenicolous species of the
genus, namely P. santessonii, confirming that it belongs to the
genus Protothelenella. Protothelenella santessonii is the only
species of the genus likely to be confined to the genus Clado-
nia. It is characterized by black perithecia, subcylindrical asci
and hyaline, submuriform ascospores often with an apiculus
(Mayrhofer 1987, Zhurbenko & Alstrup 2004). The phylogenetic
position of the genus Protothelenella and the family Protothele-
nellaceae remains uncertain within the Ostropomycetidae.
Schmitt et al. (2005) found that this family was basal to the
order Ostropales, but could not fit it in any order. In the recent
phylogeny of the Lecanoromycetes (Miadlikowska et al. 2014)
no member of the family was included. Several phylogenetic
studies have found that Protothelenella forms a well-supported
clade with Anzina (Lumbsch et al. 2007, 2012, Aptroot et al.
2014, Resl et al. 2015), a result similar to what we found here.
The genus Lettauia was first placed in the family Fuscidiaceae
(Hawksworth & Santesson 1990) on the basis of the ascus type,
similar to that of the genus Ropalospora. However, our results
placed Lettauia cladoniicola, the type species of the genus,
in the genus Cryptodiscus, family Stictidaceae, rejecting the
hypothesis that Lettauia belonged to the family Fuscidiaceae
(Table 3). So far the family Stictidaceae comprised fungi with
saprophytic, lichenized and lichenicolous lifestyles charac-
terized by a crystalline ascoma margin and long, filiform
ascospores (Wedin et al. 2005, Baloch et al. 2009, 2013).
Lettauia cladonii cola differs from the genus Cryptodiscus
basically by its non-urceolate apothecia, although also C. pini
presents superficial apothecia (Baloch et al. 2009) and by its
lichenicolous lifestyle. However, the presence of a more or
less hyaline proper exciple without embedded crystals, the
absence of periphysoids and the comparatively short, few-
celled ascospores are consistent with the genus Cryptodiscus
(Baloch et al. 2009). Therefore we propose to combine Lettauia
cladoniicola in Cryptodiscus.
The phylogenetic analyses unequivocally support that the two
newly described species, C. epicladonia and C. galaninae
belong to the genus Cryptodiscus. Morphologically, C. epi-
cladonia differs from Cryptodiscus in the presence of more
or less superficial ascomata with a crystalline rim, asci with a
narrow internal apical beak, a K/I– hymenium and asci and a
lichenicolous lifestyle. This species slightly resembles the genus
Nanostictis, a small genus of lichenicolous fungi whose hosts
mostly belong to the order Peltigerales (Christiansen 1954,
Etayo 2002, Etayo & Sancho 2008). Cryptodiscus, however, dif-
fers from Nanostictis species in several ascomatal characters.
The monophyly and phylogenetic relationship of Nanostictis
within the family Stictidaceae remain unstudied. Cryptodiscus
galaninae fits well the current concept of Cryptodiscus (Baloch
et al. 2009) except for the lichenicolous lifestyle. The placement
of these three species in the genus Cryptodiscus broadens the
generic concept presented by Baloch et al. (2009). Another
lichenicolous fungus from Stictidaceae that grows on Cladonia
is Stictis cladoniae. We have revised the type material of this
species and confirmed that it is morphologically very different
from the other species inhabiting Cladonia (see above). Sev-
eral authors doubted that this species belongs to the genus
Stictis (Christiansen 1954, Sherwood 1977), however, no fresh
material was available to solve this doubt by means of mo-
lecular data.
The genus Corticifraga was described by Hawksworth & San-
tesson (1990) as an obligately lichenicolous genus growing
on species of Peltigerales, with C. peltigerae as type species.
Currently, the genus comprises seven species and is character-
ized by initially immersed almost perithecioid or lens-shaped,
finally apothecioid ascomata, an often rather reduced exciple,
paraphyses with gradually thicked or capitate apices, clavate,
non-amyloid, 8-spored asci, and ellipsoid, soleiform, fusiform or
subcylindrical, transseptate ascospores (Hawksworth & Santes-
son 1990, Zhurbenko 2007, Etayo & Sancho 2008, Spribille et
al. 2010). Hawksworth & Santensson (1990) suggested that this
genus could belong to the order Ostropales because of the pres-
ence of nonamyloid asci. The phylogenetic analyses showed
that C. peltigerae belongs to the family Graphidaceae subfamily
Gomphilloidae, closely related to Actinoplaca strigulacea. The
species included in Gomphilloidae have rounded to elongate,
immersed to sessile apothecia, anastomosed paraphyses, non
amyloid asci, ascospores with transversal to muriform septa
and a special kind of conidiomata called hyphopores (Vězda
& Poelt 1987, Lücking et al. 2004). It is noteworthy that anas-
tomosed paraphyses and hyphopores (important characters of
Gomphilloidae) have never been observed in species of Cortici-
fraga. The current circumscription of this subfamily includes 23
genera (Rivas-Plata et al. 2012), most of which are lichenized
and live in tropical areas (Vězda & Poelt 1987, Lücking et al.
2004). However, it also includes species with a lichenicolous
lifestyle, such as Gyalideopsis cochlearifera, G. epithallina,
G. floridae, G. parvula, G. stereocaulicola and Aulaxina ag-
gregata (Lücking 1997, Lücking & Sérusiaux 1998, Etayo &
Diederich 2001, Lücking & Kalb 2002, Etayo 2010).
The coelomycetous genus Lichenosticta currently comprises
five lichenicolous species (Hawksworth 1981, Lawrey & Die
derich 2016). It is characterized by uniloculate, subglobose
to broadly pyriform, translucent brown to black, erumpent
pycnidia; branched conidiophores; enteroblastic, phialidic, acro-
pleurogenous conidiogenous cells integrated into chains; and
108 Persoonia – Volume 39, 2017
hyaline, aseptate, smooth-walled conidia (Hawksworth 1981).
Its relationship with Lecanorales was previously suggested,
since similar catenate conidiogenous cells and an entero-
blastic conidiogenesis had been found in lichenized species
(Hawksworth 1981, Vobis & Hawksworth 1981). In this study,
its phylogenetic placement in Lecanorales is confirmed by
molecular data. However, our analyses do not clarify to which
family this genus belongs, because its relation with Gypsoplaca
macrophylla was not supported. With regard to morphological
similarities, the genus Gypsoplaca has branched conidiophores
(Timdal 1990), such as those found in Lichenosticta, but the
production of conidia is always apical, while in Lichenosticta it
is both lateral and terminal.
The lichenicolous coelomycetous genus Epicladonia includes
four species (Hawksworth 1981, Ihlen & Wedin 2005), three of
which have been included in the study. This genus was resolved
as polyphyletic, forming two clades, one of which is exclusively
constituted by the type species E. sandstedei and another
formed by the other two monophyletic species, E. simplex and
E. stenospora. Epicladonia simplex and E. stenospora were
placed in the family Pilocarpaceae and E. sandstedei was
placed in the class Leotiomycetes. The polyphyly of the genus
Epicladonia is hardly surprising, since the studies based on
molecular data have proved that many anamorphic fungi, for
example Phoma (Lawrey et al. 2012), are polyphyletic. On the
other hand, it is unexpected for E. sandstedei to be phylogeneti-
cally so far from E. simplex and E. stenospora. Furthermore,
there are very few anamorphic fungi known in the class Leotio-
mycetes (Wang et al. 2006), although several genera of hypho-
mycetes have recently been placed in it (Campbell et al. 2006,
Réblová et al. 2011). Epicladonia simplex and E. steno spora
seldom induce the formation of galls and their conidia are almost
always aseptate, while E. sandstedei usually induces galls
and its conidia generally have one septum (Hawksworth 1981,
Zhurbenko & PinoBodas 2017). The family Pilocarpaceae is
mostly formed by lichenized fungi, although some species of
the genus Micarea are lichenicolous (Coppins 2009, Van den
Boom & Ertz 2014). The pycnidia of some species, such as
Fellhanera gyrophorica which has a gaping ostiole (Sérusiaux
et al. 2001), are similar to the pycnidia of Epicladonia (Hawk-
sworth 1981). The conidiogenous cells of the genus Micarea
are ampulliform to cylindrical, similar to those of Epicladonia.
However, their conidiogenesis is enteroblastic (Coppins 1983),
while in Epicladonia it is holoblastic (Hawksworth 1981).
The genus Scutula is closely related to Bacidia (Fig. 1), a result
already found by Andersen & Ekman (2005). Scutula epiblas-
tematica was related to the clade formed by S. miliaris and
S. tuberculosa. These three species together with S. heeri and
S. dedicata form Scutula s.str. (Wedin et al. 2007). Several au-
thors have pointed out that Scutula is heterogeneous (Santes-
son 1960, Triebel et al. 1997, Hawksworth 2003, Wedin et al.
2007) and they agreed on the necessity of a revision. According
to Triebel et al. (1997) and Wedin et al. (2007), Scutula s.str.
is distinguished by its lichenicolous lifestyle, lecideine apothe-
cia, an 8-spored asci with amyloid tholus and a diffuse non-
amyloid axial body, hyaline, mainly 1-septate, smooth-walled
ascospores and mitospores of different types. One species of
this genus, S. cladoniicola, has been described living on spe-
cies of Cladonia. We have studied the isotype of this species
(see the description above) and according to our observations
the reactions of asci with I and K/I are neither suggestive of
Scutula nor of Dactylospora, therefore this species may belong
to a different genus. However, we have not obtained any fresh
material to test its phylogenetic position.
Several lichenicolous fungi, so far unclassified in any class
of Ascomycota (Bachmanniomyces uncialicola, Epicladonia
stenospora, E. sandstedei, E. simplex, Epigloea soleiformis,
Lichenosticta alcicorniaria) have been placed within Lecano-
romycetes in this study. The phylogenetic positions of other
lichenicolous fungi have been confirmed or sharpened (Corti-
cifraga peltigerae, Dactylospora deminuta, D. glaucomarioides,
D. parasitica, Protothelenella santessonii and Sarcogyne sphae-
rospora). Our results offer a new approach to the family
Stictidaceae, extending the generic concept of Cryptodiscus,
which now includes species with a lichenicolous life-style.
Nevertheless, additional sampling will be necessary in order
to understand the evolution of the lichenicolous lifestyle in this
class. On the basis of the morphological characters it has been
maintained that the genera Aabaarnia, Biazrovia, Caliciella,
Catillaria, Corticiruptor, Endohyalina, Epilichen, Nimisiostella,
Normanogalla, Paralethariicola, Piccolia, Raesaenenia, Sco-
liciosporum, Spirographa, Umbilithecium and Umushamyces
belong to the Lecanoromycetes (Lawrey & Diederich 2016), but
there are no molecular studies yet that confirm this assertion.
As we have found here, more anamorphic lichenicolous fungi
might belong to Lecanoromycetes.
Acknowledgements We thank the curators of K and S herbaria for the
loans of type specimens of Scutula cladoniicola and Stictis cladoniae
and Dr. J. Kocourková (Prague), Prof. T. Ahti, A. Launis (Helsinki) and
M. Kocakaya (Yozgat) for providing specimens. Dr. Paul Diederich is cordially
thanked for critical reading the manuscript. This study has received funding
from the European Union’s Seventh Framework Programme for research,
technological development and demonstration under grant agreement no.
PIEF-GA-2013-625653 ‘CLADOF’. R. Pino-Bodas thanks the MINECO for
a Juan de la Cierva-Incorporación no. 2015-23526 support. Study of M.P.
Zhurbenko was carried out within the framework of the research project of the
Komarov Botanical Institute Russian Academy of Sciences (St. Petersburg,
Russia) no. 01201255602 and financially supported by the grant of the Rus-
sian Foundation for Basic Research no. 14-04-01031 ‘Lichenicolous fungi
of Northwest Caucasus’.
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Absconditella sphagnorum 1 AFTOLID 2315 AY300824 – AY300872 –
Absconditella sphagnorum 2 M24 EU940095 – EU940247 JX298897
Acarospora laqueata AFTOLID 1007 AY640943 AY640984 DQ991757 DQ842014
Acarosporina microspora AFTOLID 78 AY584643 AY584667 AY584612 DQ782834
Actinoplaca strigulacea AFTOLID 106 DQ782905 DQ782878 – –
Adelolecia pilati Ekman 3373 AY300826 – AY300874 –
Alectoria ochroleuca AFTOLID 209 DQ986801 DQ983483 DQ986785 HQ650597
Anaptychia palmulata AFTOLID 648 DQ883801 DQ883792 DQ912286 HQ650702
Anzina carneonivea Palize 4168 AY212829 – AY212851 AF274077
Arctomia delicatula Palice s.n. (F) AY853355 – AY853307 –
Arthrorhaphis citrinella AFTOLID 2341 AY853356 – AY853308 –
Aspicilia caesiocinerea AFTOLID 653 DQ986778 DQ986736 DQ986892 HQ650636
Aspicilia cinerea AFTOLID 647 DQ986779 DQ986735 DQ986890 HQ650637
Bacidia schweinitzii AFTOLID 642 DQ782911 DQ782884 DQ972998 DQ782850
Bacidina arnoldiana AFTOLID 1845 DQ986798 DQ986702 DQ986810 HQ650650
Baeomyces placophyllus AFTOLID 347 AF356658 AF356657 AY584695 –
Bellemerea alpina Buschbom 23.8.2000-22 AY532982 AY456692 – –
Biatora alaskana G. Thor 24732 – – KF662405 KF650958
Biatora subduplex AFTOLID 4912 KJ766533 KJ766693 KJ766360 –
Botryotinia fuckeliana AFTOLID 59 AY544651 AY544695 AY544732 DQ491491
Brigantiaea fuscolutea Gaya 65 JQ301544 JQ301604 JQ301478 –
Bryoria trichodes AFTOLID 872 DQ986752 DQ986740 DQ986896 HQ650610
Byssoloma subdiscordans Tonsberg 25968 – – AY567779 –
Calenia monospora Lücking 032h KF833327 – KF833339 –
Calopadia foliicola Lücking 16011 – – AY567782 –
Caloplaca arnoldii Gaya 5 JQ301547 JQ301606 JQ301481 JQ301657
Caloplaca chalybaea Gaya 38 JQ301550 JQ301607 JQ301484 JQ301659
Caloplaca chilensis Gaya 68 JQ301551 JQ301608 JQ301485 JQ301660
Caloplaca cinnamomea Gaya 24 JQ301552 JQ301609 JQ301487 –
Caloplaca gloriae Gaya 59 JQ301555 JQ301613 JQ301491 –
Caloplaca saxicola Soechting 7451 AJ535282 AJ535269 – –
Caloplaca scoriophila Gaya 47 JQ301560 JQ301617 JQ301496 JQ301664
Caloplaca scotoplaca Gaya 40 JQ301561 JQ301618 JQ301497 JQ301665
Candelariella reflexa AFTOLID 1271 DQ912331 DQ912331 DQ912272 –
Carbonea supersparsa AFTOLID 3696 – – – –
Carbonea vitellinaria R. Tuerk 32321 – – – AY541239
Carbonea vorticosa Tuerk 44642 – – – JN873871
Carbonicola anthracophila Timdal 11027 KF360456 – KF360424 KF360379
Carestiella socia Gilenstam 2437a AY661682 – AY661678 AY661682
Catolechia wahlenbergii 1 AFTOLID 1667 KJ766542 KJ766697 KJ766370 –
Catolechia wahlenbergii 2 AFTOLID 1743 DQ986794 DQ986704 DQ986811 HQ650649
Cecidonia umbonella Buschbom 21.08.2001-9b AY532990 – – –
Cecidonia xenophana Buschbom 26.08.2001-9 AY532991 – – –
Cetraria islandica AFTOLID 211 DQ912334 DQ912311 DQ912277 JQ301699
Chlorociboria aeruginosa AFTOLID 151 AY544669 AY544713 AY544734 DQ491501
Cladonia caroliniana AFTOLID 3 AY584640 AY584664 AY584614 DQ782832
Cladonia stipitata AFTOLID 1657 DQ973003 DQ973026 DQ972975 –
Coccocarpia domingensis AFTOLID 122 DQ912346 DQ912323 – –
Coccocarpia erythroxyli AFTOLID 333 DQ883800 DQ883791 DQ912294 HQ650691
Coccocarpia palmicola AFTOLID 1636 KJ766545 KJ766700 KJ766375 –
Coenogonium luteum AFTOLID 352 AF279387 AF279386 AY584699 HQ650710
Collema cristatum AFTOLID 1013 DQ917408 DQ917410 DQ917409 –
Crocynia pyxinoides AFTOLID 111 AY584653 AY584677 AY584615 –
Cryptodiscus foveolaris EB88 FJ904671 – FJ904693 –
Cryptodiscus gloeocapsa 1 AFTOLID 2367 AF465440 AF465456 AY300880 –
Cryptodiscus gloeocapsa 2 EB93 FJ904674 – FJ904696 –
Cryptodiscus pallidus 1 EB152 FJ904679 – FJ904701 FJ904679
Cryptodiscus pallidus 2 EB173 – – – FJ904680
Cryptodiscus pini EB178 FJ904683 – FJ904705 FJ904683
Cryptodiscus tabularum 1 Baloch SW073 FJ904688 – FJ904710 –
Cryptodiscus tabularum 2 EB77 FJ904687 – FJ904709 FJ904687
Cudoniella clavus AFTOLID 166 DQ470944 DQ470992 FJ713604 DQ491502
Dactylina arctica AFTOLID 225 DQ986802 HQ650598 DQ986786 HQ650598
Dactylospora haliotrepha AFTOLID 758 FJ176855 FJ176802 KJ766382 –
Dactylospora lobariella AFTOLID 2137 FJ176891 FJ176837 – –
Dactylospora mangrovei AFTOLID 2108 FJ176890 FJ176836 KJ766382 –
Dactylospora vrijmoediae NTOU4002 – – – KJ958534
Degelia plumbea AFTOLID 990 DQ912347 DQ912324 DQ912299 –
Dermea acerina AFTOLID 941 DQ247801 DQ247809 DQ976373 –
Diploschistes cinereocaesius AFTOLID 328 DQ883799 DQ883790 DQ912306 HQ650715
Diploschistes euganeus DNA6795 KF688507 – KF688507 KF688485
Diploschistes muscorum SFB 3 KC167077 KC167077 KC167055 KC167004
Erioderma verruculosum AFTOLID 337 DQ973041 DQ973017 DQ972990 –
Evernia prunastri AFTOLID 1272 KJ766557 KJ766713 KJ766389 HQ650611
Fellhanera bouteillei AFTOLID 4917 KJ766559 KJ766716 KJ766392 –
Fissurina insidiosa AFTOLID 1662 DQ973045 DQ973022 DQ972995 –
Fissurina sp. AFTOLID 2101 KJ766560 KJ766717 KJ766393 –
Flavocetraria nivalis AFTOLID 231 DQ883795 DQ883786 DQ912278 –
Appendix 1 List of sequences downloaded from GenBank.
Taxa ID LSU rDNA SSU rDNA mtSSU ITS rDNA
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Flavoparmelia caperata AFTOLID 2 AY584639 AY584663 AY584617 HQ650680
Fuscidea austera AFTOLID 1671 KJ766562 KJ766719 KJ766395 –
Fuscidea cyathoides AFTOLID 1672 KJ766563 – KJ766396 –
Fuscopannaria ignobilis AFTOLID 1011 DQ917417 DQ986708 DQ917416 HQ650673
Geoglossum nigritum AFTOLID 56 AY544650 AY544694 AY544740 DQ491490
Graphis scripta AFTOLID 2091 KJ440899 AF038878 KJ440959 –
Gyalecta jenensis AFTOLID 361 AF279391 AF279390 AY584705 HQ650712
Gyalidea hyalinescens AFTOLID 332 DQ973046 DQ973023 DQ972996 –
Gypsoplaca macrophylla 1 AFTOLID 1703 – KJ766722 – –
Gypsoplaca macrophylla 2 AFTOLID 3810 – KJ766722 – –
Hymenelia epulotica 1 AFTOLID 1829 KJ766569 KJ766405 KJ766405 –
Hymenelia epulotica 2 AFTOLID 1844 KJ766569 KJ766404 KJ766404 –
Hypocenomyce scalaris AFTOLID 687 DQ782914 DQ782886 DQ912274 DQ782852
Hypogymnia physodes AFTOLID 1966 JQ301600 JQ301651 JQ301541 JQ301700
Hypotrachyna degelii AFTOLID 324 DQ912337 DQ912314 DQ912281 –
Icmadophila ericetorum AFTOLID 875 DQ883694 DQ883704 DQ986897 –
Immersaria usbekica Roux 1.09.2000.5 AY532985 – – –
Imshaugia aleurites AFTOLID 1044 DQ986753 JQ301652 DQ986864 HQ650612
Ingvariella bispora BCNLich 17183 HQ659185 – HQ659174 –
Lasallia papulosa AFTOLID 650 DQ883691 DQ883701 DQ986891 HQ650603
Lasallia pustulata AFTOLID 554 DQ883690 DQ883700 DQ986889 HM161456
Lecania cyrtella AFTOLID 1791 KJ766577 KJ766732 KJ766412 HQ650645
Lecanora achariana AFTOLID 1693 DQ973027 DQ973004 DQ972976 –
Lecanora conizaeoides AFTOLID 1858 – KJ766736 KJ766418 –
Lecanora contractula AFTOLID 877 DQ986746 DQ986741 DQ986898 HQ650604
Lecanora hybocarpa AFTOLID 639 DQ782910 DQ782883 DQ912273 DQ782849
Lecanora strobilina AFTOLID 1794 KJ766583 KJ766739 KJ766420 –
Lecidea auriculata Lay 07-0075 HQ660536 HQ660520 GU074500 –
Lecidea fuscoatra AFTOLID 589 DQ912332 DQ912310 DQ912275 HQ650707
Lecidea laboriosa AFTOLID 1388 KJ766586 DQ986727 DQ986882 –
Lecidea silacea AFTOLID 1368 – DQ986723 DQ986878 HQ650629
Lecidella elaeochroma AFTOLID 1275 DQ986747 – – HQ650605
Lecidoma demissum AFTOLID 1376 DQ986759 DQ986726 DQ986881 HQ650630
Leotia lubrica AFTOLID 1 AY544644 AY544687 AY544746 DQ491484
Lepraria lobificans AFTOLID 325 DQ986768 DQ986733 DQ986887 –
Leptogium lichenoides AFTOLID 1015 DQ917412 DQ917413 DQ923120 HQ650672
Letrouitia domingensis Gaya 55 JQ301569 JQ301625 JQ301505 –
Letrouitia vulpina Gaya 72 JQ301571 JQ301627 JQ301509 –
Lithographa tesserata P95 KJ462346 KR017261 KR017340 KJ462269
Lobaria scrobiculata AFTOLID 128 AY584655 AY584679 AY584621 –
Lobariella palliola AFTOLID 310 DQ883796 DQ883787 DQ912296 HQ650695
Lobothallia radiosa AFTOLID 1860 KJ766596 KJ766746 KJ766430 –
Lopezaria versicolor AFTOLID 108 DQ912353 DQ912330 AY584622 –
Maronea chilensis AFTOLID 370 AY640955 AY640994 KJ766432 –
Massjukiella candelaria AFTOLID 4377 JQ301587 JQ301639 JQ301528 –
Megalospora sulphurata Gaya 73 JQ301573 – JQ301514 –
Megalospora tuberculosa AFTOLID 107 AY584650 AY584674 AY584623 HQ650701
Melanelia fuliginosa AFTOLID 1370 DQ986803 DQ983485 DQ986787 HQ650599
Micarea alabastrites Andersen 17 AY756327 – AY567764 AY756469
Micarea denigrata AFTOLID 4923 KJ766598 KJ766750 KJ766437 –
Miriquidica garovaglii 1 Szczepanska 538 KF562180 – KR995350 KF562188
Miriquidica garovaglii 2 AFTOLID 2688 – – AY567711 –
Mollisia cinerea AFTOLID 76 DQ470942 DQ470990 DQ976372 DQ491498
Mycobilimbia lurida AFTOLID 1859 KJ766653 KJ766789 KJ766486 –
Mycobilimbia tetramera AFTOLID 1637 KJ766600 – KJ766439 –
Mycoblastus sanguinarius AFTOLID 196 DQ912333 DQ782879 DQ912276 DQ782842
Myriotrema olivaceum Lumbsch 19113f & Mangold EU075627 – EU075579 –
Nephroma parile AFTOLID 131 AY584656 AY584680 AY584625 HQ650698
Nesolechia oxyspora 1 Wedin 7890 GU994613 – GU994659 GU994568
Nesolechia oxyspora 2 Ertz 16840 (BR) KR995417 – – KR995295
Ochrolechia trochophora AFTOLID 880 KJ766609 DQ986743 DQ986901 –
Ochrolechia yasudae AFTOLID 882 DQ986776 DQ986744 DQ986902 –
Ophioparma lapponica AFTOLID 1707 DQ973028 DQ973005 DQ972977 –
Ophioparma ventosa AFTOLID 1694 KJ766610 – KJ766447 –
Orceolina kerguelensis AFTOLID 296 AF274116 DQ366257 AY212830 –
Ostropa barbara EB85 – – – HM244773
Parmelina tiliacea AFTOLID 1307 KJ766616 KJ766759 KJ766451 –
Parmotrema tinctorum AFTOLID 7 AY584635 AY584659 AY584627 HQ650684
Peltigera degenii AFTOLID 134 AY584657 AY584681 AY584628 –
Peltigera sp. AFTOLID 1838 DQ986796 DQ986705 DQ986809 HQ650648
Peltula auriculata AFTOLID 892 DQ832330 DQ832332 DQ922953 DQ832329
Peltula umbilicata AFTOLID 891 DQ832334 DQ782887 DQ922954 DQ832333
Pertusaria amara AFTOLID 1067 AF274101 AF274104 AY584713 HQ650677
Pertusaria hemisphaerica AFTOLID 959 AF381556 DQ902340 DQ973000 HQ650676
Petractis clausa Hafellner A 1 AF356662 AF356661 – –
Petractis nodispora AFTOLID 7804 FJ588713 FJ588712 – –
Phacopsis vulpina D132 – – – AF450285
Phaeophyscia orbicularis AFTOLID 1308 DQ912343 DQ912320 DQ912289 JQ301694
Appendix 1 (cont.)
Taxa ID LSU rDNA SSU rDNA mtSSU ITS rDNA
114 Persoonia – Volume 39, 2017
Phaeopyxis punctum 1 TU65586 KJ559567 KJ559587 – –
Phaeopyxis punctum 2 TU68298 KJ559568 KJ559588 – –
Phaeopyxis punctum 3 Diederich 17303 – KJ559591 – KJ559551
Phlyctis argena AFTOLID 1375 – – DQ986880 –
Phyllobaeis erythrella AFTOLID 329 DQ986780 DQ986734 DQ986888 –
Phyllobaeis imbricata AFTOLID 852 DQ986781 DQ986739 DQ986895 HQ650635
Physcia aipolia AFTOLID 84 DQ782904 DQ782876 DQ912290 DQ782836
Physconia muscigena AFTOLID 220 DQ912344 DQ912321 DQ912291 JQ301696
Placynthiella oligotropha AFTOLID 1797 – KJ766766 KJ766458 –
Platismatia glauca AFTOLID 201 DQ973032 DQ973007 DQ972980 –
Platythecium grammitis AFTOLID 2095 KJ766627 KJ766769 KJ766461 –
Pleopsidium chlorophanum AFTOLID 1004 DQ842017 DQ525541 DQ991756 –
Pleopsidium gobiense AFTOLID 1003 DQ883698 DQ525573 DQ991755 HQ650723
Polychidium muscicola AFTOLID 230 DQ986770 DQ986731 DQ986885 HQ650626
Polysporina arenacea SAR275 LN810814 – LN810939 LN810814
Polysporina subfuscescens 1 CR26058 – – KM879329 KM879334
Polysporina subfuscescens 2 CR26059 – – KM879330 KM879333
Porina lectissima Arup & Baloch SW152 HM244774 – HM244756 –
Porpidia albocaerulescens AFTOLID 1246 DQ986757 DQ986716 DQ986871 –
Porpidia speirea AFTOLID 1050 DQ986758 DQ986711 DQ986865 DQ986711
Protoblastenia calva AFTOLID 992 JQ301601 JQ301653 DQ986904 HQ650618
Protoblastenia rupestris AFTOLID 4911 KJ766631 KJ766771 – –
Protopannaria pezizoides AFTOLID 222 DQ912350 DQ912326 DQ912301 HQ650693
Protoparmelia atriseda Ponzetti 26046 KF562182 – – KF562190
Protoparmelia cupreobadia Fryday 863 KF562184 – – KF562192
Protoparmelia phaeonesos Timdal 11000 KF562185 – – KF562193
Protothelenella corrosa Palice 2002 AY607734 – AY607746 –
Protothelenella sphinctrinoidella Lumbsch 19031d AY607735 – AY607747 –
Pseudephebe pubescens AFTOLID 1775 KJ766635 KJ766773 KJ766467 –
Pseudocyphellaria anomala AFTOLID 132 DQ883794 DQ883785 DQ912298 HQ650697
Psilolechia leprosa Tonsberg & Botnen 27362 AY756333 – AY567730 AY756496
Psora decipiens AFTOLID 4857 KJ766640 KJ766778 KJ766474 –
Punctelia rudecta AFTOLID 9 AY584636 AY584660 AY584630 HQ650686
Puttea margaritella M149 EU940038 EU940111 EU940261 EU940187
Pycnothelia papillaria AFTOLID 1377 DQ986800 DQ983481 DQ986783 HQ650595
Pyxine subcinerea AFTOLID 686 DQ883802 DQ883793 DQ912292 HQ650705
Raesenenia huuskonenii Myllys 040811-53 KR995426 AF450289 – KR995306
Ramalina complanata AFTOLID 966 DQ883783 DQ883784 DQ972986 HQ650720
Ramalina farinacea AFTOLID 1965 KJ766646 KJ766783 KJ766480 –
Ramboldia elabens AFTOLID 4996 KJ766648 KJ766784 KJ766482 –
Ramboldia gowardiana AFTOLID 4913 KJ766649 KJ766785 – –
Ramboldia insidiosa AFTOLID 1756 KJ766650 KJ766786 – –
Rhizocarpon oederi AFTOLID 1372 DQ986804 DQ983486 DQ986788 –
Rhizoplaca melanophthalma AFTOLID 2383/2384 DQ787351 – DQ787352 –
Rimularia limborina isolate 1062 KJ462349 KR017277 KJ462404 KJ462273
Rinodina tephraspis AFTOLID 1314 DQ912345 DQ912322 DQ912293 –
Ropalospora chlorantha AFTOLID 884 – – KJ766487 –
Sagiolechia protuberans AFTOLID 7896 KJ766655 – HM244757 –
Sarcogyne algoviae SAR37 LN810849 – LN810975 LN810849
Sarcogyne clavus SAR220 LN810853 – – LN810853
Sarcogyne hypophaea Pykala 23561 LN810857 – – LN810857
Sarcogyne plicata AFTOLID 4830 KJ766657 KJ766791 – –
Sarcogyne regularis AFTOLID 3292 – – AY853343 –
Schizoxylon albescens 1 AFTOLID 4193 – – DQ401142 –
Schizoxylon albescens 2 Gilenstam 2696a DQ401144 – – HQ287353
Sclerococcum sphaerale 1 Diederich 17283 JX081673 – JX081678 –
Sclerococcum sphaerale 2 Ertz 17425 JX081674 – JX081676 –
Scoliciosporum intrusum Ekman s. n. AY756329 – AY567767 –
Scutula krempelhuberi Wedin 6356 – – AY567789 –
Scutula miliaris Wedin 6850 – – AY567790 –
Solenopsora candicans AFTOLID 1277 KJ766660 KJ766795 KJ766493 –
Spaerophorus fragilis AFTOLID 226 DQ986805 DQ983487 DQ986805 HQ650600
Sphaeropezia capreae 1 EB-2010 – – HM244751 –
Sphaeropezia capreae 2 HM244772 AY661684 – AY661674 –
Sphaeropezia mycoblasti EB-2012b JX266159 – JX266157 –
Sphaeropezia sp. 2 EB-2012c JX266160 – – –
Sphaeropezia yckselensis EB-2012a JX266159 – JX266156 –
Sphaerophorus globosus AFTOLID 1057 DQ986767 DQ986712 DQ986866 HQ650622
Stictis confusum Wedin 7070 DQ401143 – DQ401141 DQ401143
Stictis populorum Gilenstam 2610a AY527327 – AY527356 AY527327
Stictis radiata AFTOLID 398 AF356663 U20610 AY584727 DQ782846
Stictis urceolatum AFTOLID 96 DQ986790 – DQ986790 HQ650601
Strangospora pinicola AFTOLID 4980 KJ766664 KJ766803 KJ766500 –
Teloschistes exilis AFTOLID 87 AY584647 AY584671 FJ772245 –
Teloschistes flavicans AFTOLID 315 JQ301578 JQ301631 JQ301520 JQ301685
Tephromela atra AFTOLID 780 – DQ986737 DQ986894 HQ650606
Appendix 1 (cont.)
Taxa ID LSU rDNA SSU rDNA mtSSU ITS rDNA
115
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
Tetramelas phaeophysciae 1 Nordin 4922 – – – DQ198359
Tetramelas phaeophysciae 2 Nordin 5663 – – – DQ201951
Tetramelas pulverulentus 1 Nordin 4417 – – – DQ201952
Tetramelas pulverulentus 2 Nordin 4427 – – – DQ201953
Thamnolia vermicularis AFTOLID 2071 AY853395 AF085472 AY853345 –
Thelenella antarctica Lumbsch 19006a AY607739 – AY607749 –
Thelotrema lepadinum AFTOLID 83 – – DQ972997 HQ650717
Thrombium epigaeum Lumbsch 11179 AY607741 – AY607751 –
Trapelia placodioides AFTOLID 962 AF274103 AF119500 AF431962 –
Trapeliopsis flexuosa AFTOLID 1825 KJ766668 KJ766807 KJ766505 –
Trichoglossum hirsutum AFTOLID 64 AY544653 AY544697 AY544758 DQ491494
Umbilicaria aprina AFTOLID 1416 DQ986799 DQ986706 DQ986814 HM161480
Umbilicaria arctica AFTOLID 1266 DQ986772 DQ986717 DQ986872 HM161454
Umbilicaria muelhenbergii AFTOLID 404 AY640977 AY641016 AY584729 –
Umbilicaria spodochroa AFTOLID 555 DQ986773 DQ986707 DQ986815 HM161481
Usnea antarctica AFTOLID 813 DQ883692 DQ883702 DQ990920 HQ650616
Vulpicida pinastri AFTOLID 198 DQ912285 – – –
Wawea fruticulosa AFTOLID 3401 DQ007347 – DQ871023 –
Xanthomendoza fallax Gaya 33 JQ301580 JQ301633 JQ301522 –
Xanthomendoza poeltii Gaya 7 JQ301583 JQ301636 JQ301525 –
Xanthoparmelia conspersa AFTOLID 4 AY584641 AY584665 AY584633 HQ650688
Xanthoria aureola Gaya 9 JQ301585 JQ301637 JQ301526 –
Xanthoria elegans AFTOLID 214 DQ912352 DQ912329 DQ912304 –
Xanthoria parietina Gaya 8 JQ301589 JQ301641 JQ301530 –
Xanthoria polycarpa AFTOLID 200 DQ912351 DQ912328 DQ912303 –
Xylographa parallela AFTOLID 4895 KJ766679 – KJ766516 –
Xyloschistes platytropa AFTOLID 4891 KJ766680 – KJ766517 –
Appendix 1 (cont.)
Taxa ID LSU rDNA SSU rDNA mtSSU ITS rDNA
116 Persoonia – Volume 39, 2017
ITS rDNA LSU rDNA SSU rDNA mtSSU
Code Best BLAST hit % Simila Evalue Best BLAST hit % Simila Evalue Best BLAST hit % Simila Evalue Best BLAST hit % Simila Evalue
rity/bp rity/bp rity/bp rity/bp
RP43 Phaeopyxis punctum, 98 % /525 0.0 – – – – – – – – –
P. punctum KJ559545
RP68 Uncultured fungus, 86 %/370 2e-102 Micarea adnata, 83 %/445 3e-136 – – – – – –
E. stenospora KC965887 AY756326
RP93 Phaeopyxis punctum, 95 %/525 0.0 Phaeopyxis punctum, 100 %/ 850 0.0 – – – – – –
P. punctum KJ559545 KJ559568
RP94 Phaeopyxis punctum, 92 % /529 0.0 – – – – – – – – –
P. punctum KJ559551
RP95 Phaeopyxis punctum, 95 %/509 0.0 Phaeopyxis punctum, 99 %/ 738 0.0 – – – – – –
P. punctum KJ559551 KJ559567
RP96 Phaeopyxis punctum, 100 %/525 0.0 Phaeopyxis punctum, 99 %/ 996 0.0 – – – – – –
P. punctum KJ559545 KJ559568
RP97 Phaeopyxis punctum, 97 % /528 0.0 – – – Phaeopyxis punctum, 99 % /924 0.0 – – –
P. punctum KJ559551 KJ559588
RP106 Uncultured fungus, 99 %/498 0.0 Fungal sp., 98 %/ 355 2e-175 Helotiales sp., 99 % /308 1e-157 Leotiomycetes sp., 99 %/782 0.0
E. sandstedei KF617768 KT289722 LN901162 KT263275
RP109 Rhizoplaca macleanii, 83 %/ 468 6e-128 – – – – – – – – –
L. alcicorniaria JX036152
RP119 Squamarina gypsacea 87 %/324 1e-93 Psilolechia leprosa, 90 % /911 0.0 – – – Micarea micrococca, 93 %/ 762 0.0
E. stenospora AY756333 EF453683
RP123 Phaeopyxis punctum, 93 %/ 529 0.0 – – – – – – – – –
B. uncialicola KJ559551
RP127 Uncultured fungus, 98 %/478 0.0 – – – – – – – – –
D. ahtii KC965673
RP159 Uncultured Cryptodiscus, 89 %/520 1e-174 Bryophagus gloeocapsa, 95 % /773 0.0 Teloschistes flavicans, 96 %/398 0.0 Bryophagus gloeocapsa , 90 %/638 0.0
L. cladoniicola KP323396 AF465440 JQ301631 AY300880
RP160 Uncultured Cryptodiscus, 89 %/520 1e-174 Bryophagus gloeocapsa, 95 % /773 0.0 Bryophagus gloeocapsa, 98 %/ 349 7e-174 Bryophagus gloeocapsa, 90 %/741 0.0
L. cladoniicola KP323396 AF465440 AF465456 AY300880
RP168 Rhizoplaca macleanii, 84 %/542 2e-133 – – – Gypsoplaca macrophylla, 98 %/936 0.0 Lecanora hybocarpa, 90 % /723 0.0
L. cladoniicola JX036152 KJ66722 EF105417
RP182 Uncultured fungus, 98 %/477 0.0 – – – – – – Porina lucida, 85 %/100 1e-23
D. ahtii KC965673 FJ11132
RP189 Uncultured fungus, 86 %/370 1e-105 Palicella glaucopa, 81 %/592 3e-162 – – – – – –
E. stenospora KC965887 KJ152458
RP190 Lecidella aff. euphorea, 83 %/ 363 1e-100 Pisolechia leprosa, 90 %/ 737 0.0 Micarea adnata, 96 % /1538 0.0 – – –
E. stenospora KT453756 AY756333 AF455134
RP203 – – – – – – – – – Anzina carneonivea, 88 % /561 0.0
E. soleiformis AY212851
RP204 Uncultured fungus, 92 %/196 4e-68 Xylographa hians, 84 %/ 846 0.0 – – – Thrombium epigaeum, 83 %/ 590 4e-167
E. soleiformis KF617618 KJ462359 AY607750
Appendix 2 Results of BLAST searches for each new sequences generated in this study, bp represent the covertage in pairs of bases.
117
R. Pino-Bodas et al.: Cladoniicolous fungi placed within Lecanoromycetes
RP205 Uncultured soil fungus, 89 % /523 3e-180 Prototelenella sphinc- 97 %/898 0.0 Rhizoplaca chrysoleuca, 94 % /740 0.0 Protothelenella corrosa, 98 % /780 0.0
P. santessonii KC965473 trinoidella, AY607735 AY530883 AY607746
RP206 Uncultured soil fungus, 89 % /523 3e-180 Prototelenella sphinc- 97 %/895 0.0 Rhizoplaca chrysoleuca, 94 % /668 0.0 – – –
P. santessonii KC965473 trinoidella, AY607735 AY530883
RP208 Uncultured fungus, 92 %/508 0.0 – – – – – – Cryptodiscus faveolaris, 88 %/ 664 0.0
C. epicladonia KF617267 AY661673
RP23 Uncultured fungus, 98 %/477 0.0 Uncultured fungus, 87 % /557 0.0 – – – Dactylospora mangrovei, 88 %/549 0.0
D. ahtii KC965673 KP889692 KJ766383
RP235 Uncultured fungus, 99 %/460 0.0 – – – – – – Chaetothryales, 85 % /502 2e-159
D. deminuta KC966342 KT263240
RP263 Fungal endophyte, 93 % /304 0.0 – – – – – – Leotiomycetes sp., 96 %/ 499 0.0
E. sandstedei HQ335298 KT263275
RP275 Uncultured fungus, 85 %/467 1e-119 Dactylospora mangrovei, 95 %/949 0.0 – – – Dactylospora mangrovei, 88 %/617 0.0
D. glaucomarioides KC965719 FJ176890 KJ766383
RP276 Bacidia circumspecta, 94 %/482 0.0 – – – – – – Scutula krempelhuberi, 91 %/ 165 5e-60
S. epiblastematica AF282124 AY567789
RP282 Uncultured soil fungus, 87 % /380 9e-112 Monilinia fructicola, 83 % /714 0.0 – – – Calenia monospora, 85 % /583 0.0
C. peltigerae GU211937 AY544683 KF833339
RP301 Uncultured soil fungus, 87 % /380 9e-112 Acarospora umbilicata, 96 %/ 1031 0.0 – – – Polysporina subfuscescens, 99 %/739 0.0
S. sphaerospora GU211937 LN810808 LN10967
RP314 Cryptodiscus pini, 89 %/ 523 4e-174 – – – Xylographa vitiligo, 97 % /865 0.0 – – –
C. galaninae FJ904682 AY779284
RP352 Phaeopyxis punctum, 97 %/528 0.0 – – – Phaeopyxis punctum, 99 % /1482 0.0 – – –
B. uncialicola KJ559551 KJ55959
RP362 – – – Psilolechia leprosa, 92 % /943 0.0 Micarea adnata, 96 % /1534 0.0 – – –
E. stenospora AY756333 AF455134
RP391 – – – Dactylospora mangrovei, 94 % /926 0.0 – – – Dactylospora mangrovei, 91 %/591 0.0
Dactylospora sp. FJ176890 KJ766383
RP392 – – – Biatora subduplex, 96 %/ 1078 0.0 – – –
E. stenospora KJ766693
RP395 Rhizoplaca macleanii, 83 %/545 5e-129 Porpidia glaucophaea, 93 %/975 0.0 – – – – – –
L. cladoniicola JX036152 AY532950
RP422 Ascomycete sp., 95 %/480 0.0 Ascomycete sp., 97 %/ 1028 0.0 – – – Dactylospora mangrovei, 95 %/656 0.0
D. parasitica EF210107 EF210108 KJ766383
RP423 – – – – – – – – – Dactylospora mangrovei, 90 %/604 0.0
D. parasitica KJ766383
RP424 – – – Ascomycete sp., 97 %/ 538 0.0 – – – Dactylospora mangrovei, 95 %/614 0.0
D. parasitica EF210108 KJ766383
RP426 Biatora alaskana, 87 % /304 6e-88 – – – – – – – – –
E. simplex KF650958
RP427 Biatora alaskana, 87 % /304 1e-89 – – – – – – – – –
E. simplex KF650958
RP428 Biatora alaskana, 84 % /304 3e-71 – – – – – – – – –
E. simplex KF650958
