Four new species of Bacidia s.s. (Ramalinaceae, Lecanorales) in the Russian Far East

Abstract

The molecular phylogeny of Bacidia s.s. in the Russian Far East was investigated using 62 nucleotide sequences from the ITS nrDNA region, 22 of which were newly obtained. Phylogenetic reconstructions employed Bayesian inference and maximum likelihood searches using MrBayes and RAxML. In addition, ITS2 secondary structures added further support using Compensatory Base Changes. As a result of morphological and phylogenetic studies, four new species of Bacidia are described. Bacidia areolata sp. nov. belongs to the suffusa group. It was collected once in Khabarovskiy Krai, the Russian Far East, on the bark of Acer tegmentosum and is closely related to B. suffusa but differs in having a smooth, cracked to areolate thallus and shorter spores. Bacidia elongata sp. nov. is a member of the fraxinea group and is similar to B. fraxinea but differs in having a wide zone of cells with enlarged lumina along the edge of the exciple. In fact, this zone of enlarged cells, in combination with its overall habit, places it morphologically close to B. suffusa , B. millegrana and B. campalea . Bacidia kurilensis sp. nov. is a basal member of the laurocerasi group and closely related to B. biatorina , B. heterochroa , B. laurocerasi and B. salazarensis. However, the combination of a granular thallus, large black apothecia and a green hue in the upper part of the exciple edge as well as in the epihymenium sets it apart from the species mentioned above. Bacidia sachalinensis sp. nov. resolves as a strongly supported member of the polychroa group and is known from a single locality in Sakhalin, the Russian Far East. Its thallus structure and apothecium colour are variable, which is typical for the polychroa group, but it differs from B. polychroa by having shorter spores with fewer septa and a mainly smooth to areolate thallus.

Full text

The Lichenologist 50(6): 603–625 (2018) © British Lichen Society, 2018
doi:10.1017/S0024282918000397
Four new species of Bacidia s.s. (Ramalinaceae,Lecanorales)
in the Russian Far East
Julia V. GERASIMOVA, Aleksandr K. EZHKIN and Andreas BECK
Abstract: The molecular phylogeny of Bacidia s.s. in the Russian Far East was investigated using 62
nucleotide sequences from the ITS nrDNA region, 22 of which were newly obtained. Phylogenetic
reconstructions employed Bayesian inference and maximum likelihood searches using MrBayes and
RAxML. In addition, ITS2 secondary structures added further support using Compensatory Base
Changes. As a result of morphological and phylogenetic studies, four new species of Bacidia are
described. Bacidia areolata sp. nov. belongs to the suffusa group. It was collected once in Khabarovskiy
Krai, the Russian Far East, on the bark of Acer tegmentosum and is closely related to B. suffusa but differs
in having a smooth, cracked to areolate thallus and shorter spores. Bacidia elongata sp. nov. is a member
of the fraxinea group and is similar to B. fraxinea but differs in having a wide zone of cells with enlarged
lumina along the edge of the exciple. In fact, this zone of enlarged cells, in combination with its overall
habit, places it morphologically close to B. suffusa,B. millegrana and B. campalea.Bacidia kurilensis sp.
nov. is a basal member of the laurocerasi group and closely related to B. biatorina,B. heterochroa,
B. laurocerasi and B. salazarensis. However, the combination of a granular thallus, large black apothecia
and a green hue in the upper part of the exciple edge as well as in the epihymenium sets it apart from the
species mentioned above. Bacidia sachalinensis sp. nov. resolves as a strongly supported member of the
polychroa group and is known from a single locality in Sakhalin, the Russian Far East. Its thallus
structure and apothecium colour are variable, which is typical for the polychroa group, but it differs from
B. polychroa by having shorter spores with fewer septa and a mainly smooth to areolate thallus.
Key words: Bacidiaceae, compensatory base changes, crustose lichens, diversity, ITS secondary
structure, morphology, phylogenetic analysis
Accepted for publication 28 March 2018
Introduction
The lichen flora of the Russian Far East has
been investigated for more than a century
(Brummitt 2001). In particular, Primorskiy
Krai, the most south-eastern region of Russia,
is comparatively well studied (Skirina &
Moiseyevskaya 2004 and references therein).
However, large areas and many lichen genera
remain largely unexplored. One genus we
know little about in this region is Bacidia.
Previous reports of Bacidia from this area can
be found in a number of lichen checklists and
papers (Brotherus et al. 1936; Skirina 1995,
2015; Tchabanenko 2002; Galanina 2008;
Kuznetsova et al. 2013).
Historically, Bacidia included crustose
lichens with a chlorococcoid photobiont and
biatorine apothecia producing ascospores
that have at least three septa (Zahlbruckner
1921–1940). During the second half of the
20th century, Bacidia sensu Zahlbruckner was
partially split up and numerous species were
transferred to other taxa, resulting in the
recognition of more than 20 genera (e.g.
Santesson 1952; Vězda 1978, 1986, 1991;
Hafellner 1984; Lücking 1992; Sérusiaux
1993; Ekman 1996).
The first phylogenetic study (Ekman
2001) indicated that many species referred to
J. V. Gerasimova: Komarov Botanical Institute of the
Russian Academy of Sciences, Prof. Popova St. 2,
197376 St. Petersburg, Russia. Email: jgerasimova.
lich@yandex.ru
J. V. Gerasimova and A. Beck: SNSB-BSM, Bota-
nische Staatssammlung München, Menzinger Straße
67, D-80638 München, Germany.
A. K. Ezhkin: Institute of Sea Geology and Geophysics,
Far East Branch of the Russian Academy of Sciences,
Nauki St. 1B, 693022 Yuzhno-Sakhalinsk, Russia.
A. Beck: GeoBio-Center, Ludwigs-Maximilians-
Universität München, Richard-Wagner-Str. 10,
D-80333 München, Germany.
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as Bacidia, in particular those with blue-
green pigmentation in the epithecium and/or
with fusiform or bacilliform spores, were
more closely related to Toninia s.l. Bacidia in
the strict sense is consequently characterized
by acicular spores and a well-developed,
prosoplectenchymatic proper exciple com-
posed of radiating, abundantly furcate and
rarely anastomosed hyphae with heavily
gelatinized cell walls and cell lumina that
become compressed and narrower with age.
Since this first phylogenetic study, a limited
set of Bacidia s.s. species have been investi-
gated further (Andersen & Ekman 2005;
James et al. 2006; Reese Næsborg et al. 2007;
Jeon et al. 2009; Schmull et al. 2011; Sér-
usiaux et al. 2012; Miadlikowska et al. 2014;
Mark et al. 2016; Lendemer et al. 2016).
We present here a taxonomic study of
Bacidia s.s. using morphological and mole-
cular data and focusing on material from the
Russian Far East. In this area, members of
the genus are abundant in habitats that have
high humidity and moderate insolation.
These include open forests, forest edges,
swamps, river banks and valleys, as well as hill
and mountain slopes close to the sea or near
lakes or swamps. The aim of the present study
was to clarify species boundaries in members
of Bacidia s.s. from the Russian Far East.
Material and Methods
The material for this study primarily consisted of 83 fresh
collections of Bacidia gathered in the field by the authors
in the southern part of the Russian Far East (Primorskiy
and Khabarovskiy Krai), as well as on Sakhalin and the
Kurile Islands, from 2013 to 2015. All material was
collected in old forest communities ranging from about
sea level up to 1350 m. The forests were mainly in
floodplains and were mixed conifer-broadleaf and
spruce-fir forests in river valleys which were open with a
high humidity. Herbarium material from the Pacific
Institute of Geography (PIG, without acronym) and the
Botanical Garden-Institute of FEB RAS (VBGI) was
also studied. In order to confirm the taxonomic position
of Bacidia suffusa (Fr.) A. Schneid. from the Russian Far
East and verify the status of a specimen of that species
from GenBank (AF282091), two additional specimens
of B. suffusa and one specimen of B. diffracta S. Ekman
that had been collected in North America were studied
and sequenced. The type material of all species present
in this study was analyzed. Voucher specimens
were deposited in the herbaria of the Botanische
Staatssammlung München (M) and Komarov Botanical
Institute RAS (LE). Detailed information of the newly
obtained sequences together with their respective vou-
cher information and GenBank Accession numbers are
given in Table 1. GenBank Accession numbers addi-
tionally included in the phylogenetic analyses are given
in Supplementary Table S1 (available online). Voucher
information for all investigated specimens are given in
Supplementary Table S2 (available online). Localities
and herbarium numbers for all specimens investigated in
the course of this study are given in the Supplementary
Material (available online). As Japan is geographically
close, we included species of Bacidia recorded in
checklists of the lichens of Japan (Kashiwadani & Inoue
1993; Inoue 1994; Harada et al. 2004; see Table 2).
Bacidia arceutina (Ach.) Arnold, B. laurocerasi (Delise ex
Duby) Zahlbr., B. polychroa (Th. Fr.) Körb., B. rosella
(Pers.) De Not., B. rubella (Hoffm.) A. Massal., B.
schweinitzii (Fr.) A. Schneid. and B. subincompta (Nyl.)
Arnold have been recorded before and not specifically
from Japan and are therefore not included in Table 2.
Morphology
All specimens were examined using standard micro-
scopic techniques following Ekman (1996). Microscopic
observations were made using light microscopes and a
Zeiss Axioplan microscope equipped with differential
interference contrast. Micrographs of cross-sections
were taken on a Zeiss Axio Imager microscope with an
attached AxioCam MRc5 camera and processed with the
Zeiss ZEN2012 image program. Images of the external
features of the species were obtained from a Zeiss Stemi-
2000 CS microscope with an attached AxioCam MRc5
camera and processed with the Zeiss AxioVision image
program. The photomicrographs with detail of exciple
structures in Fig. 4C were taken with a Zeiss AxioScope
A1 compound microscope equipped with a Canon 60D
digital camera. Spore and apothecium measurements are
given as mean ðxÞ±1SD with outliers in parentheses. All
other measurements are given as minimum, mean ðxÞ
and maximum values. In those cases where not enough
measurements were available for the calculation of a
mean value, only minimum and maximum values
are given.
DNA extraction, PCR amplification and DNA
sequencing
DNA extraction was carried out using PCR Template
Preparation Kit (Roche, Mannheim, Germany) follow-
ing the manufacturer’s instructions. 5–8 apothecia were
used from fresh material not older than 3 years and
thallus fragments were removed in order to minimize the
risk of contamination.
PCR amplifications for the ITS1, ITS2 and 5.8S
regions were performed using 5 µl 5× Green GoTaq
®
Flexi buffer, 1·75 µl MgCl
2
,2·5µl dNTPs, 1·25 µl of each
primer, 0·1µl Taq polymerase and 1–5µl of DNA solu-
tion in 25 µl volume. Cycling conditions included initial
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denaturation at 95 °C for 2 min, 5 cycles of 95 °C for 45
s, 54 °C for 60 s and 72 °C for 60 s, 33 cycles of 95 °C for
45 s, 52 °C for 60 s and 72 °C for 60 s, with a final
extension step at 72 °C for 7 min. We used the primers
ITS1F (White et al. 1990) and ITS4m as described in
Beck & Mayr (2012) or, for old herbarium specimens,
primers ITS3 and ITS4 (White et al. 1990).
All PCR products were run on an agarose gel, cut out
under UV-light and purified with the PCR Clean-Up &
Gel Extraction Kit (SLG, Gauting, Germany). Purified
products were labelled with the BigDye Terminator
v3.1 Kit (Applied Biosystems, Darmstadt, Germany).
Cycle sequencing consisted of 30 cycles of 95 °C for
10 s, 50 °C for 15 s and 60 °C for 3 min, using the PCR
primers individually. Post-sequencing clean-up was
performed using gel filtration with Sephadex G-50
Superfine (GE Healthcare, Uppsala, Sweden) following
the manufacturer’s protocol. Forward and reverse
strand sequences were detected on an ABI 3730 48-
capillary automatic sequencer (Applied Biosystems)
and assembled using the program PhyDE (http://www.
phyde.de/index.html).
Alignment and phylogenetic analyses
BLAST searches in GenBank were performed to detect
and exclude accessory/lichenicolous fungi and potential
contaminants. Alignments were carried out using stan-
dard settings in MUSCLE v.3.8.31 (Edgar 2004) as
implemented in the program PhyDE and optimized by
hand using the ITS2 secondary structures (see below) as a
guide. Positions that possessed numerous indels and pre-
sented a nucleotide in less than 3% of the sequences as
well as ambiguously aligned regions were excluded.
Two datasets were analyzed for this study. Using the
first dataset, we aimed to ensure that the new sequences
belonged to Bacidia s.s. and to examine the relationships
of this genus within the broader context of the Ramali-
naceae. This large dataset comprised 130 sequences that
included: 1) the sequences of all representatives of for-
mer Bacidiaceae used in the publication by Ekman
(2001); 2) all other ITS sequences of Bacidia as well as
Bacidina available from GenBank (Groner & LaGreca
1997; James et al. 2006; Reese Næsborg et al. 2007; Jeon
et al. 2009; Schmull et al. 2011; Czarnota & Guzow-
Krzemińska 2012; Sérusiaux et al. 2012; Miadlikowska
et al. 2014; Lendemer et al. 2016; Mark et al. 2016); 3)
the ITS sequences of Bacidia s.s. generated in this study.
The second dataset was restricted to Bacidia s.s.
sequences, based on the results of the first analysis. This
allowed the use of a larger part of the alignment because
it did not contain ambiguous parts.
The ITS nrDNA sequence dataset was subjected to
maximum likelihood (ML) and Bayesian inference (BI)
analyses. To select the nucleotide substitution model
and parameters for the ML searches, a statistical selec-
tion of best-fit models was carried out in jModelTest
2.1.5 (Guindon & Gascuel 2003; Darriba et al. 2012).
TABLE 1. Specimens used in the phylogenetic study of Bacidia together with their voucher information and GenBank Accession
numbers.
Specimen Locality Voucher
ITS GenBank
Accession number
Bacidia areolata Russia, Far East Gerasimova M-0182592 (M) MH048614
B. circumspecta Russia, Far East Gerasimova L-13006 (LE) MH539764
B. diffracta USA, Minnesota Wetmore 46555-A (M) MH048620
B. elongata Russia, Far East Gerasimova M-0182571 (M) MH048626
B. elongata Russia, Far East Gerasimova M-0182625 (M) MH048627
B. elongata Russia, Far East Gerasimova M-0182626 (M) MH048628
B. elongata Russia, Far East Gerasimova M-0182627 (M) MH048629
B. friesiana Russia, Far East Gerasimova L-13159 (LE) MH539765
B. kurilensis Russia, Far East Ezhkin M-0182620 (M) MH048610
B. kurilensis Russia, Far East Ezhkin M-0182621 (M) MH048611
B. kurilensis Russia, Far East Ezhkin M-0182622 (M) MH048612
B. laurocerasi Russia, Far East Galanina 424 (VBGI) MH048609
B. rubella Russia, Far East Gerasimova M-0182581 (M) MH048630
B. sachalinensis Russia, Far East Ezhkin M-0182619 (M) MH048621
B. sachalinensis Russia, Far East Ezhkin M-0182623 (M) MH048622
B. sachalinensis Russia, Far East Ezhkin SAK 147 (SAK) MH048623
B. sachalinensis Russia, Far East Ezhkin SAK 148 (SAK) MH048624
B. sachalinensis Russia, Far East Ezhkin M-0182624 (M) MH048625
B. schweinitzii Russia, Far East Gerasimova M-0182580 (M) MH048613
B. suffusa USA, Louisiana Tucker 17000 (M) MH048618
B. suffusa USA, Minnesota Wetmore 40219 (M) MH048619
B. suffusa Russia, Far East Gerasimova M-0182593 (M) MH048616
B. suffusa Russia, Far East Gerasimova M-0182594 (M) MH048617
B. suffusa Russia, Far East Gerasimova M-0182601 (M) MH048615
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The optimal model was identical using either the cor-
rected Akaike information criterion or the Bayesian
information criterion (TIM2ef + I + G). The model
provided estimations of equal nucleotide frequencies, a
rate matrix with six different substitution types, assum-
ing a heterogeneous rate of substitutions with a gamma
distribution of variable sites (number of rate cate-
gories =4, shape parameter a =0·7160) and a proportion
of invariable sites (pinvar) of 0·2781. Heuristic phyloge-
netic searches were conducted using 100 random addi-
tion sequence (RAS) replicates, tree bisection-
reconnection branch swapping (TBR), saving all trees
and collapsing branches with a maximum length equal to
zero using PAUP* v4.0b10 (Swofford 2002). Due to the
size of the dataset, the most likely phylogeny was calcu-
lated using PAUP* as this allows the use of a more
extensive search option than for instance RAxML, which
computes only an approximate log likelihood score of the
alternative topology after subtree reinsertion (Stamatakis
et al. 2008). However, support values were calculated in
independent runs using RAxML and MrBayes to allow
better comparability with other studies. Support values
using a bootstrap search in PAUP* are indicated on the
respective branches of the phylogeny.
TABLE 2. Bacidia species from Japan studied in this research for comparison with the newly described species from
the Russian Far East.
Name according to Japanese
checklist* Comments Reference
Bacidia abducens (Nyl.) Zahlbr. Currently: Bacidia schweinitzii (Fr. ex Tuck.)
A. Schneid.
Ekman 1996: 99
B. akagiensis (Vain.) Yasuda Apothecia blackish; spores fusiform-oblong,
16–24 × 4–5µm, with 3 septa
Vainio 1921: 66
B. baculifera (Nyl.) Zahlbr. Thallus whitish; apothecia convex, c.0·5mm
diam.; spores bacilliform, 32–42 × 5–6µm,
with 1–7 septa
Nylander 1890: 67
B. beckhausii Körb. Currently: Biatora beckhausii (Körb.) Tuck. Printzen 2014: 451
B. endoleucula (Nyl.) Zahlbr. Currently: Bacidia laurocerasi (Delise ex Duby)
Zahlbr.
Ekman 1996: 82
B. hakkodensis Kashiw. Apothecia pale brown to yellowish brown; spores
oblong-ellipsoid, 27–35 × 5–6µm, with
3–5(–7) septa
Kashiwadani & Sasaki 1987: 69
B. hakonensis (Müll. Arg.) Yasuda Apothecia black; spores obovate-cylindrical,
20–35 × 7–10 µm, with 3–5 septa
Müller 1892: 198
B. invertens (Nyl.) Zahlbr. Currently: Bacidia laurocerasi (Delise ex Duby)
Zahlbr.
Ekman 1996: 82
B. leptoboliza (Nyl.) Zahlbr. Belongs to Lecanactis Printzen 1995: 190
B. luteorufula (Tuck.) Zahlbr. Apothecia yellowish to orange; spores unicellular,
ovoid, fusiform to ellipsoid, 5·0×2·5µm
Tuckerman 1866 [1864]: 276
B. micrommata (Kremp.) R. Sant. Currently: Eugeniella micrommata (Kremp.)
Lücking, Sérus. & Kalb.
Lücking 2008: 716
B. myricicola (Vain.) Yasuda Probably belongs to Phyllopsora According to observation of
S. Ekman [TUR-V 20326]
B. spumosula (Zahlbr. ex Yasuda)
Yasuda
Apothecia black; spores unicellular, ellipsoid
to oval, 11–13 µm×7–8µm
Yasuda 1925: 28
“Bacidia subcontes (Nyl.) Anzi”Name doesn’t exist; probably misspelling of
Bacidia subincompta
B. subdiscendens (Nyl.) Zahlbr. Apothecia dark brown to blackish; hypothecium
brown-black; spores acicular, 55–65 × 3 µm,
with 7–9 septa
Nylander 1890: 67
B. subrudis (Nyl.) Zahlbr. Apothecia blackish; epithecium yellow-brownish;
spores oblong, 25–34 × 0·8–1·0µm, with 3–7
septa
Nylander 1890: 64
B. subvermifera (Nyl.) Zahlbr. Apothecia black; hypothecium blackish; spores
vermiform, 30–40 × 3 µm, with 3–5 septa
Nylander 1900: 33
B. uvulina Zahlbr. Apothecia black; spores vermiform, 27–36 × 1·6–
3·5µm, with uneven (up to 7) septa
Zahlbruckner 1916: 52
B. yasudae (Vain.) Yasuda Belongs to Micarea According to observation of
S. Ekman [TUR-V 20783]
*Kashiwadani & Inoue 1993; Inoue 1993; Harada et al. 2004.
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Further ML analysis was performed using RAxML
v8.2.4 on both datasets using 1000 rapid bootstrap
pseudo-replicates, following a GTRGAMMA model of
molecular evolution (Stamatakis 2014). Bayesian infer-
ence was carried out using the Markov chain Monte
Carlo method (MCMC) using MrBayes v3.2.6 (Ron-
quist et al. 2012). As the model recommended by jMo-
delTest (TIM2ef + I + G) is not available in MrBayes,
the GTRGAMMA model was selected instead based on
the recommendation of Huelsenbeck & Rannala (2004).
Two parallel runs were performed (two cold chains) with
a single tree saved every 100th generation for a total of
10 000 000 generations. As convergence was reached
after 25% of the trees, the initial 25% was discarded as
burn-in and the results summarized as a 50% majority-
rule consensus tree.
The phylogenetic trees were visualized using FigTree
v1.4.2 (Rambaut 2009). Only clades that received
bootstrap support ≥70% in ML and PP ≥0·95 in BI were
considered highly supported.
ITS secondary structures
As additional evidence, we further analyzed
ITS2 secondary structure, using compensatory base
changes (CBCs) and hemi-CBCs in the structurally con-
served regions of helix III (Coleman 2003). CBCs are
mutations that occur in a primary RNA transcript, whereby
both nucleotides paired in the secondary configuration of
the ITS transcript mutate so that their bond is retained (e.g.
G-C mutates to A-U). A hemi-CBC (hCBC) is the muta-
tion of one of the two nucleotides while maintaining the
nucleotide bond. Most likely ITS2 secondary structures of
the RNA transcript were determined by delimiting the
highly conserved start and end region of the first three
helices. The structure of these sequence sections was
deduced using the RNAfold web server (http://rna.tbi.
univie.ac.at/cgi-bin/RNAfold.cgi), folding helices I, II, III
and IV individually and therefore more reliably. In all cases
we used the minimum free energy (MFE) structure
obtained. The depictions of ITS2 secondary structure were
made for morphologically close species of Bacidia s.s.,
corresponding to the species groups as indicated on the
phylogenetic tree. Comparisons among sequences from
each group are shown, using the entire ITS2 structure of
one species per group as a “core”and indicating nucleotide
substitutions on this core. The following sequences have
been used as core: Bacidia rubella (GenBank, AF281087)
for Bacidia fraxinea group, B. laurocerasi (AF282080), B.
polychroa (AF282089), and B. suffusa (MH048616) for
respective groups.
Results
Morphology
Ten species of Bacidia s.s. were dis-
tinguished based on morphological analysis,
including the previously described Bacidia
friesiana (Hepp) Körb., B. rubella (Hoffm.)
A. Massal., B. laurocerasi (Delise ex Duby)
Zahlbr., B. polychroa (Th. Fr.) Körb.,
B. schweinitzii (Fr.) A. Schneid. and B.
suffusa (Fr.) A. Schneid. As no name was
available for four morphologically distinct
entities, B. areolata,B. elongata,B. kurilensis
and B. sachalinensis are described as new. In
addition, B. schweinitzii is reported from
Russia for the first time.
The revision of all available herbarium
collections (172) resulted in a new determi-
nation for some specimens. A specimen of
B. arceutina (Ach.) Arnold, previously reported
from a single locality in Kamchatka (Nesha-
taeva et al. 2004), was identified as B. laur-
ocerasi. Specimens from our own collections,
initially described as B. fraxinea Lönnr., were
finally determined to be B. elongata sp. nov.
One herbarium specimen (PIG 28798), pre-
viously labelled as B. fraxinea and which was
old with a damaged thallus, had several apo-
thecia with only a few spores with uneven
septa. We were unable to differentiate this
species from the closely related B. rubella,
consequently its identification is uncertain.
The herbarium specimens of B. biatorina
(Körb.) Vain. (e.g. Galanina 2008; Skirina
2015) were redetermined as B. schweinitzii,
B. friesiana or as belonging to Bacidia s.l.
Therefore, the occurrence of B. biatorina in
the Russian Far East also remains question-
able. The specimen of B. rosella (Pers.) De
Not., previously reported from a single
locality (Yakovchenko et al. 2013), is related
to B. suffusa. In conclusion, there are four
species, B. arceutina,B. biatorina,B. fraxinea
and B. rosella, which should probably not be
included in the list of Bacidia s.s. that occur
in the Russian Far East.
Detailed morphological examination was
carried out for previously sequenced speci-
mens belonging to B. diffracta,B. polychroa
and B. suffusa (GenBank AF282090,
AF282089, AF282091) and these were
compared with specimens from the Russian
Far East.
Based on morphological studies, it should
be noted that the new species are character-
ized by variations in thallus structure and
apothecium colour, which is characteristic of
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the whole genus (Ekman 1996). Features of
Far Eastern species are summarized in
Table 3 including, for comparison, all mor-
phologically similar species.
Phylogeny
The first alignment of the larger dataset
comprised 130 sequences and 485 char-
acters. Tylothallia biformigera was selected as
outgroup. The new specimens of Bacidia
from the Russian Far East were shown to
belong to Bacidia s.s. However, three speci-
mens of Bacidia sp. from GenBank
(KX098339, KX098340 and KX098341)
were found to belong to the Bacidina group
and were therefore not included in the sub-
sequent analyses.
The second alignment of the reduced
dataset contained 62 sequences (including
22 obtained for this study) and 481 char-
acters, with Bacidia incompta,Biatora globu-
losa,B. hemipolia and Cliostomum griffithii as
outgroups. This dataset contained 20
Operational Taxonomic Units (OTUs) of
Bacidia s.s.
The ML and BI analyses recovered highly
concordant topologies of the phylogenetic
trees. Only the phylogeny resulting from the
ML search in PAUP* is presented here as
there were no contradictions in supported
parts of the trees.
The backbone within Bacidia s.s. is poorly
resolved; only species groups were recovered
with significant support values but the rela-
tionships between these are unclear. As a
result of the phylogenetic analyses, a number
of well-supported clades can be recognized
within the Bacidia s.s. clade (Fig. 1).
Bacidia polychroa,B. diffracta and
B. sachalinensis belong to a highly supported
polychroa group (ML/BI: 98/1·0) and split
into two main clades, with B. sachalinensis as
sister to the other species. Two sequences of
B. diffracta specimens group together with
GenBank sequences of B. suffusa from the
USA (AF282091) in a supported clade (92/
0·99). Given that B. sachalinensis is recovered
as a strongly supported group, as well as its
morphological differences, it is described
here as a species new to science.
Sister to the polychroa group is a group
including B. elongata,B. fraxinea and
B. rubella (74/0·99). The last two form a clade
(82/0·98), with the position of B. rubella
remaining uncertain. In addition, as only one
sequence of B. fraxinea was included in the
analysis, we consider this group needs further
work using a larger number of samples.
Specimens of B. elongata sp. nov. form a
monophyletic group with strong support
(100/0·99).
Bacidia laurocerasi,B. biatorina and
B. kurilensis form a strongly supported clade
(100/1·0), the laurocerasi group (Fig. 1).
Within this group, another strongly sup-
ported clade (100/1·0) includes B. laurocerasi
sequences from the Russian Far East
(MH048609) and North America (GenBank
AF282078). A second clade within this
group contains B. biatorina, and a third is
composed of three sequences of B. kurilensis
sp. nov. with high support (76/1·0). Bacidia
biatorina was placed as sister to B. kurilensis in
the heuristic search using PAUP*, but with-
out support. In contrast, B. biatorina is sister
to B. laurocerasi in RAxML and BI with
strong support (95/1·0). In all analyses the
relationships within the schweinitzii group are
only weakly supported, require further
investigation and are not discussed here.
Bacidia suffusa and B. areolata form a
strongly supported group (98/1·0). Sequen-
ces of B. suffusa were split into two main
lineages: Far Eastern populations (100/1·0)
and those from North America (98/0·96).
The sequence representing B. areolata sp.
nov. was placed as sister to B. suffusa, form-
ing a separate branch and described here as
new to science.
There are sequences of several other spe-
cies of Bacidia available which were included
in the phylogeny but which were not found
in the Russian Far East. An example is
B. absistens (Nyl.) Arnold which is so far
known from a single locality in European
Russia (Gerasimova 2016). Bacidia lutescens
Malme and B. hostheleoides (Nyl.) Zahlbr.
belong to a well-supported clade with long
branches and both of which are widely dis-
tributed in the Neotropics (Malme 1935;
Ekman 1996). Species of the highly
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AF282076_Cliostomum griffithii
76/0.98/94
98/1.0/- 100/1.0/96
72/0.93/-
92/0.99/-
98/1.0/-
100/1.0/100
97/1.0/99 76/0.97/-
74/0.99/-
82/0.98/-
78/0.98/-
100/1.0/95
100/1.0/99
100/1.0/82
100/1.0/-
100/1.0/-
97/1.0/-
100/1.0/74
76/1.0/-
98/1.0/91
0.03
100/1.0/87
99/1.0/96
98/0.96/87
97/1.0/95
100/0.99/99
AF282073_Biatora globulosa
AF282072_Biatora hemipolia
AF282092_Bacidia incompta
AF282083_Bacidia arceutina
AF282084_Bacidia scopulicola
AF282085_Bacidia absistens
AF282091_Bacidia suffusa
AF282090_Bacidia diffracta
AF282089_Bacidia polychroa
MH048620_Bacidia diffracta
MH048621_Bacidia sachalinensis
MH048622_Bacidia sachalinensis
MH048623_Bacidia sachalinensis
MH048624_Bacidia sachalinensis
MH048625_Bacidia sachalinensis
MH048626_Bacidia elongata
MH048627_Bacidia elongata
MH048628_Bacidia elongata
MH048629_Bacidia elongata
AF282088_Bacidia fraxinea
MH048630_Bacidia rubella
EU266078_Bacidia rubella
KX132984_Bacidia rubella
JQ796852_Bacidia rubella
HQ650644_Bacidia rubella
AF282087_Bacidia rubella
AF282082_Bacidia lutescens
AF282081_Bacidia hostheleoides
AF282078_Bacidia laurocerasi
MH048609_Bacidia laurocerasi
MH048610_Bacidia kurilensis
MH048611_Bacidia kurilensis
MH048612_Bacidia kurilensis
MH048614_Bacidia areolata
MH048615_Bacidia suffusa
MH048616_Bacidia suffusa
MH048617_Bacidia suffusa
MH048618_Bacidia suffusa
MH048619_Bacidia suffusa
MH048613_Bacidia schweinitzi
AF282079_Bacidia biatorina
DQ782850_Bacidia schweintzii
KX151767_Bacidia sorediata
KX151769_Bacidia sorediata
KX151773_Bacidia sorediata
KX151775_Bacidia sorediata
KX151768_Bacidia sorediata
KX151770_Bacidia sorediata
KX151771_Bacidia sorediata
KX151772_Bacidia sorediata
KX151774_Bacidia sorediata
KX151761_Bacidia schweinitzii
KX151766_Bacidia schweinitzii
KX151762_Bacidia schweinitzii
KX151763_Bacidia schweinitzii
AF282080_Bacidia schweinitzii
KX151764_Bacidia schweinitzii
KX151765_Bacidia schweinitzii
AF282086_Bacidia rosella
JQ796853_Bacidia sipmanii
arceutina group
polychroa group
fraxinea group
laurocerasi group
schweinitzii group
suffusa group
JQ796851_Bacidia arceutina
FIG. 1. The most likely tree generated by phyologenetic analysis of ITS1 and ITS2 regions and 5.8S gene in
PAUP* ML analysis and representing the phylogenetic relationships of Bacidia s.s. Bootstrap support ≥70% in
RAxML analysis (first value), posterior probability ≥0·95 (second value) and ≥65% bootstrap support in ML
analysis by PAUP* (third value) were considered as highly supported and denoted by very thick lines. Support
values between 50 and 70% were considered as weakly supported and are indicated by lines of medium thickness.
New sequences are in bold.
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supported arceutina group, including B. sco-
pulicola (Nyl.) A.L. Sm. and B. sipmanii
M. Brand et al., are also not present in the
Russian Far East. As mentioned above, the
occurrence of B. rosella in the Russian Far
East also remains questionable.
ITS secondary structure
Differences in the ITS2 secondary structure
support all main groups of Bacidia s.s. and the
different lineages within the clades (Fig. 2),
often involving CBCs and hemi-CBCs in the
structurally conserved regions of helix III. By
comparison, helices I, II and IV show only
minor variation between groups and clades.
The four newly described species are sup-
ported by differences in their ITS2 secondary
structure, mainly in hemi-CBCs. Thus, by
comparison with the core (Bacidia suffusa,
GenBank MH048616), B. areolata differs
from B. suffusa and B. elongata by the
Bacidia fraxinea group Bacidia laurocerasi group
Bacidia polychroa
g
roup Bacidia suffusa group
III III
II II
IV IV
I
III III
II II
IV IV
II
I
AB
CD
FIG. 2. Differences in secondary structure of ITS2 among groups within Bacidia s.s. Variable nucleotides among
species within the groups are marked with diamonds, CBCs and hemi-CBCs are indicated by broad and narrow
arrows, respectively.
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presence of three hemi-CBCs, and from B.
fraxinea by five hemi-CBCs (Fig. 2D & A).
Bacidia sachalinensis differs from B. polychroa
by one CBC, G–U instead of C–G (Fig. 2).
Bacidia kurilensis differs from B. laurocerasi
by two CBCs and one hemi-CBC in the
conserved part of helix III (U–G instead of
C–U, and C–G instead of G–U, respectively;
Fig. 2C). Bacidia biatorina differs from B.
laurocerasi by one CBC, where a U–A pairing
was observed instead of G–C in the latter.
The ITS2 secondary structures in the
schweinitzii group correspond to the clades in
the trees. The two subclades of B. sorediata
differ by one hemi-CBC, but in the case of
the first group (including KX151767 etc.)
there is an unpaired A–G that would reveal
the presence of one CBC. Several schweinitzii
subgroups form sister clades, but without
support, while ITS2 secondary structure
supports this grouping, based on three hemi-
CBCs in helix III. These ITS secondary
structure subgroups are comprised of:
AF282080, KX151764 & KX151765;
KX151761 & KX151766; DQ782850,
KX151762 & KX151763, and a separate
lineage of MH048613 from the Russian
Far East.
The comparisons within the lutescens-
hostheleoides “group”reveal one CBC and
three hemi-CBCs in helix III as compared to
the other groups. This “group”is very diverse
with high nucleotide variation in the other
helices and many specimens have not yet
been sequenced.
Discussion
Since the first phylogenetic study, more than
50 additional ITS sequences of Bacidia s.s.
have been added to GenBank, including the
sequences obtained in this study. This has
enabled the phylogeny of Bacidia s.s. to be
refined and has made possible a better inter-
pretation in the wider context. Phylogenetic
relationships within Bacidia s.s. agree with
the previous results presented by Ekman
(2001), with the exception of the lutescens-
hostheleoides “group”which is not considered
to be a natural group. It has an uncertain
position with weak support. Both taxa have
sequences that have evolved a great variety of
nucleotides, which was demonstrated parti-
cularly in ITS2 secondary structure (see
Results). This indicates that the grouping in
this part of the phylogeny most likely results
from incomplete species sampling rather
than from a natural relationship. Thus long-
branch attraction artefacts may occur and
further sampling in this group is required to
draw reliable conclusions.
Our study indicates that much of the diver-
sity within Bacidia s.s. still needs to be investi-
gated. Of the ten species found in the Russian
FarEast,fourwerenotknownfromgeo-
graphically close areas and thus are new to
science. Table 2 provides a comparison of
Bacidia species from Japan with those newly
described species from the Russian Far East.
The additional study of ITS2 has dis-
tinguished and supported all main groups of
the genus. Despite a number of well-supported
clades being recognized, the backbone within
Bacidia s.s. remains poorly resolved. To over-
come this problem and the relationships
among the groups, sequences from additional
loci are needed for follow-up studies.
Bacidia polychroa group
All species within the polychroa group share
the K+ purplish/violet reaction in cross-
sections of apothecia. The morphology of
B. polychroa was first examined in a broad
sense, including “typical”B. polychroa,
granular B. diffracta, the specimen from
GenBank first identified as B. suffusa
(AF282091), and specimens from the Rus-
sian Far East initially identified as “Bacidia
polychroa sp.”The European specimen of
B. polychroa from GenBank (AF282089),
which was placed as a sister to B. diffracta
(Fig. 1), was a typical morph and conse-
quently corresponds to the type of B. poly-
chroa.TheB. polychroa typical morph is
characterized by a wrinkled to warted thallus
of scattered or contiguous areoles that
become finally granular. It has orange-brown
to dark red-brown apothecia with a distinct
orange-brown hypothecium which is darker
than the exciple below. Bacidia sachalinensis
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has a mainly cracked to areolate thallus and
lighter orange to orange-brown apothecia
which is also characteristic of the North
American morph of B. polychroa (see
Table 3). The separation of B. sachalinensis
from B. polychroa is also supported by hemi-
CBCs in the ITS2 secondary structure and a
low similarity of the sequences (94%).
Two species belonging to B. diffracta and a
misidentified specimen of B. suffusa from
North America (AF282091) present two
distinct morphotypes. The specimen pre-
viously referred to B. suffusa is probably a
separate species which is more closely related
to B. diffracta. However, it represents an
intermediate form between B. diffracta and
B. polychroa and is characterized by a
smooth, partly granular thallus and orange-
brown to purplish brown apothecia as well as
the typical K+ violet reaction (see Table 3).
This could mean that either the European
and North American specimens of B. poly-
chroa are different species or simply that the
lineage sorting is not complete between B.
diffracta and B. polychroa. Both representa-
tives of the granular forms of B. diffracta,
AF282090 and MH048620, are similar to
one another. The sequences reveal differ-
ences between them but, as only one speci-
men of each has been analyzed, they are
treated here as a single species.
Bacidia sachalinensis exhibits substantial
variation in thallus structure and apothecial
colour, even in the same specimen, and this is
also typical for both B. diffracta and B. poly-
chroa. To study whether B. polychroa from the
Far East was close to B. polychroa in a strict
sense, we obtained several additional sequen-
ces from Far Eastern specimens, using as
much variation in thallus and apothecial
structure as possible with the intention of
using these characters to delimit species.
Despite this variation, all specimens examined
in the phylogenetic analyses were found to
belong to B. polychroa s.s. Only a single Eur-
opean specimen of B. polychroa (Sweden) and
material from Sakhalin were included in the
current analyses. Further study is required
using collections from other parts of Europe,
the Far East and North America to reveal the
heterogeneity of this group.
Bacidia fraxinea group
Bacidia fraxinea and B. rubella represent
two morphologically well-distinguished spe-
cies, differing mainly in thallus structure
(Ekman & Nordin 1993). Our dataset is not
yet sufficient to make conclusions about the
relationship between these morphospecies.
Currently there is not enough evidence to
challenge the recognition of B. rubella and
B. fraxinea as distinct species because only
one sequence of B. fraxinea is available.
Further work is necessary here, so we
recommend retaining the morphologically
differentiated species.
Bacidia elongata represents a morphologi-
cally discrete, monophyletic entity, distinct
from B. fraxinea mainly in its exciple struc-
ture. Consequently, it is here described as a
new species. It falls close to the fraxinea
group but the zone of enlarged cells (Fig. 3C)
is characteristic for B. elongata and is rather
exceptional. There are several species of
Bacidia with a zone of enlarged cells such as
B. russeola but it is closely related to B. lauro-
cerasi (laurocerasi group) and B. heterochroa.
We suggest that this zone of enlarged cells is a
character which has evolved more than once
and B. elongata is the first example known in
the fraxinea group. These morphological
differences are also supported by the low
similarity of sequences (93–95%) and hemi-
CBCs in the ITS2 secondary structure.
Bacidia laurocerasi group
Bacidia laurocerasi from the Russian Far
East is placed as sister to the specimen for the
USA on a zero-length branch, which con-
firms its identification (Fig. 1). Furthermore,
morphological analysis has shown that
B. laurocerasi from the Far East represents the
“typical”form, corresponding to B. lauro-
cerasi subsp. laurocerasi as detailed by Ekman
(1996). This form is characterized by a
poorly defined smooth thallus, black apo-
thecia and brown epihymenium and exciple
edge. In contrast, the four sequences of the
new species B. kurilensis, sister to B. lauro-
cerasi, reveal a morphologically discrete
entity, characterized by a granular thallus and
a greenish pigment in the epihymenium and
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TABLE 3. Main diagnostic features of the Russian Far Eastern species of Bacidia s.s., including several morphologically close species for comparison. Morphologically similar groups are
indicated by grey or white banding.
Apothecium Exciple
Species of
Bacidia Thallus Colour Pruina Rim
Cell layer
along rim
Hymenium
(µm) Epithecium Hypothecium Spores (µm)
*Pigments,
K+ / −
diffracta granular brown-orange
to dark
purplish
brown
+/−brown-orange
to orange-
brown
absent 68–97 indistinctly
coloured,
colourless
to orange-
brown
pale brown-
orange to
orange-
brown
32–69 × 1·9–4·1,
with 3–11
septa
most pigmented
parts K+
purple-red
suffusa
(GenBank)
smooth to
cracked to
wrinkled
orange,
orange-
brown to
purple-
brown
+/−laterally
orange-
brown
with 1–2 cell
layers
62·5–67·5 pale orange
to orange-
brown
orange 32·5–51·0×1·7–
3·0, with 3–7
septa
most parts K+
int. or brown
part of K +
purplish
polychroa cracked to
areolate
brown-orange
to dark
purplish
brown
+/−brown-orange
to orange-
brown
without or with
single cell
layer up to
9×5µm
56–102 colourless to
orange-
brown
±brown-
orange to
dark brown
31–74 × 1·9–5·0,
with 2–15
septa
most pigmented
parts K+
purple-red
sachalinensis cracked to
areolate to
warted
pale orange to
red-brown,
rarely dark
+/−pale brown to
orange-
brown
1–2 cell layers
up to 8 × 7 µm
70·0–92·5 indistinctly
coloured,
colourless
to orange-
brown
pale brown-
yellow to
brown-
orange
36·7–63·5×2·0–
4·0, with 1–8
septa
most pigmented
parts K+
purple-red
biatorina granular orange-brown
to dark
purplish
brown
−orange-brown
to dark red-
brown
1–2 cell layers
up to
12 × 6 µm
83–87 brown-
orange to
red-brown
almost
colourless
42–57 × 2·1–2·9,
with 3–15
septa
pigmented parts
exciple and
epithecium
K±int./K+
purplish
heterochroa smooth, areolate,
rimose
purple-brown
to black
+/−brown to red-
brown
1 cell layer up
to 6 µm wide
75–115 brown colourless or
pale yellow
32–73 × 2·5–4·3,
with (3–)7–15
septa
pale yellow K+
int. or brown
parts K+
purplish
kurilensis granular to
granular isidiose
reddish brown
to almost
black
−dark brown
with greenish
hue
single cell layer
up to 6 × 5 µm
80·0–107·5 dark brown
with a dirty
green hue
almost
colourless
to pale
yellow
41–88 × 2·0–4·0,
with 3–17
septa
pigmented parts
K±int.
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TABLE 3(continued ).
Apothecium Exciple
Species of
Bacidia Thallus Colour Pruina Rim
Cell layer
along rim
Hymenium
(µm) Epithecium Hypothecium Spores (µm)
*Pigments,
K+ / −
laurocerasi smooth to
wrinkled to
warted
purple-brown
to purple-
black
−dark red-brown
to black-
brown
single cell layer
up to 9 × 6 µm
71–131 dark brown yellowish 50–108 × 1·9–
3·7, with 7–28
septa
brown parts K+
purplish
salazarensis crustose, ±rimose black +/ −red-brown
sometimes
with green in
upper part
1 cell layer up
to 6 µm wide
60–80 green colourless to
pale yellow
34–60 ×2·5–4·0,
with 5–7 septa
pale yellow
parts K+ int.
or red-brown
parts K+
purplish
areolata smooth to
areolate
pale pink to
purple-
brown
+/−yellow to
brown-
orange
3–4 cell layers
up to
12 × 6 µm
56·5–90 pale orange
to orange-
brown
pale yellow,
almost
colourless
40–82 × 2·5–4·5,
with 6–15
septa
most pigmented
parts K+ int.
elongata (dark
morph)
wrinkled to
granular
dark orange-
brown to
dark purple-
brown
+/−dark orange-
brown
4 cell layers up
to
20·0×5·5µm
62·5–110·0 pale yellow,
pale
orange,
rare
brown-
orange
pale yellow-
brown to
orange-
brown
39–80 × 2·0–4·0,
with 2–16
septa
brown parts
K+ purplish,
yellow K+
int.
suffusa smooth, wrinkled
to warted
yellow-brown
to purplish to
black
+ pale yellow to
black-brown
4–6 cell layers
up to
12 × 6 µm
77·5–137·5 yellowish pale yellow to
black-
brown
40–95 × 2·5–5·0,
with 6–17
septa
yellow to orange
parts K+ int.
elongata (pale
morph)
smooth to
areolate
orange to
orange-
brown
+/−pale orange-
brown
4 cell layers up
to
20·0×5·5µm
62·5–110·0 colourless pale yellow,
almost
colourless
39–80 × 2·0–4·0,
with 2–16
septa
K−
fraxinea smooth to
areolate
orange-brown
to
darkbrown
+/−straw to pale
orange
without or with
1 cell layer up
to 6 × 6 µm
76–103 colourless to
straw
pale orange,
straw
42–109 × 2·5–
4·3, with 3–17
septa
pigmented parts
K+ int.
*int. =intensifying
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AB
CD
EF
GH
FIG. 3. New species of Bacidia from the Russian Far East. A & B, Bacidia areolata, holotype (M-0182592). C,
B. elongata, dark morph, Primorskiy Krai (M-0182625). D, B. elongata, light morph, Khabarovskiy Krai, holotype
(M-0182571). E & F, B. kurilensis, Kurile Island, holotype (M-0182620). G & H, B. sachalinensis, Sakhalin,
holotype (M-0182619); G, smooth to warted thallus with light apothecia; H, cracked thallus with dark apothecia.
Scales: A–G=0·5 mm; H =1·0 mm. In colour online.
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upper part of the exciple (for details see
Taxonomy). These morphological differ-
ences are also supported by the low similarity
of the sequences (95%) and hemi-CBCs in
the ITS2 secondary structure. The green
pigmentation in the upper part of the hyme-
nium in B. kurilensis is similar to that of
B. heterochroa and B. salazarensis B. de Lesd.,
which also has a granular thallus. Neither B.
heterochroa nor B. salazarensis were repre-
sented in this phylogenetic study.
Bacidia schweinitzii group
This group comprises B. schweinitzii and
B. sorediata Lendemer & R.C. Harris. Spe-
cimens of the schweinitzii group were
recoveredinmultiplecladeswithinboth
B. schweinitzii s.s. (i.e. esorediate popula-
tions) and the sorediate morphotype (Baci-
dia sorediata), in accordance with the
findings of Lendemer et al. (2016). The
sequence from the Russian Far East speci-
men is placed basal to all other members of
that group, but without significant support
(59/0·93). In spite of this, according to
Lendemer et al. (2016), B. schweinitzii from
the Far East belongs to B. schweinitzii s.s. It
is characterized by a granular thallus, black
apothecia, blue-green pigmentation in the
epihymenium and a dark reddish brown
hypothecium. The specimen from GenBank
(AF282080) was also obtained from a typi-
cal B. schweinitzii with a granular thallus and
black apothecia. The internal branches in
the schweinitzii s.l. part of the tree do not
have significant support for the most part,
leaving any relationships within this group
unclear.
Bacidia suffusa group
The final group of the tree combines two
morphologically indistinct lineages of B. suf-
fusa from the Russian Far East and from
North America. Both have characters which
correspond with those of type material and
can be referred to as the typical morph
(Table 3). They probably represent different
populations or cryptic species which can be
separated geographically. Owing to the lack
of good morphological characters, they are
treated here on the population level.
Although B. areolata is represented by only
one sequence, there is quite strong morpho-
logical evidence for considering it to be a
discrete species. Moreover, this is also sup-
ported by a low similarity of the sequences
(92%) and hemi-CBCs in the
ITS2 secondary structure. There are three
hemi-CBCs between B. suffusa (Fig 2D) and
B. areolata (not shown), and these are located
in the most conserved region of helix III
(Coleman 2009).
Taxonomy
Bacidia areolata J. Gerasimova &
A. Beck sp. nov.
MycoBank No.: MB 821184
Similar to Bacidia suffusa but distinguished mainly by an
areolate thallus and lighter-coloured apothecia.
Type: Russia, Khabarovskiy Krai, Khabarovskiy
Rayon, Bolshekhekhtsirskiy State Natural Reserve, 48°
25'N, 134°77'E, 160 m, coniferous-broadleaf forest, on a
terrace above the river, on bark of Acer tegmentosum,6
September 2013, J. V. Gerasimova s. n. (M M-0182592—
holotype; LE L-13014, UPS L-721140—isotypes).
GenBank Accession no: MH048614
(Figs 3A & B, 4A & B)
Thallus poorly defined, either thin, partly
smooth to areolate, of scattered, discrete or
contiguous, flattened or ±convex areoles, or
thick, continuous, wrinkled to warted; never
with distinct granules; white, greyish green to
deep green. Prothallus present between dis-
crete areoles, white or rarely black along the
border of the thallus. Photobiont chlor-
ococcoid green alga, 6–15 × 8–18 μm.
Apothecia (0·3–)0·4–0·5–0·6(–0·9) mm,
±plane when young, remaining plane when
mature or becoming slightly convex, eprui-
nose, rarely with thin white pruina on the
edge and disc of young and medium-aged
apothecia. Disc pale pink, peach-coloured,
pale beige to pale brown, rarely purple-brown
when mature, often mottled. Margin pale
pink, yellow-brown, raised above disc in
young apothecia, later becoming ±plane.
Exciple laterally 49·0–60·4–80·0µm wide,
without crystals or sometimes with radiating
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AB
D
C
EF
GH
FIG. 4. New species of Bacidia from the Russian Far East. A & B, Bacidia areolata: A, TS apothecium; B, ascus
with spores. C & D, B. elongata: C, TS apothecium; D, spore. E & F, B. kurilensis: E, TS apothecium; F, ascus
with spores. G & H, B. sachalinensis: G, TS apothecium; H, ascus with spores. Scales: A, C & G =50 µm;
E=100 µm; B, D, F & H =20 µm. In colour online.
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clusters of crystals dissolving in N. Rim yel-
low, yellow-brown, brown-orange, often
darker in the upper part than in the lower part,
3–4 cell layers thick along the edge, distinct
zone of enlarged cells with lumina that are
7–12 µm long and 4–6µm wide; inner part
paler than or ±concolorous with rim, down-
wards almost colourless to pale yellowish, K
−.Hymenium 56·5–76·2–90·0µm high, in
lower part colourless; upper part pale orange
to orange-brown, K+ yellow. Hypothecium
pale yellow, almost colourless, K±yellow
(reaction unclear). Paraphyses simple, thin,
1·0–1·5µm wide in mid-hymenium, ±clavate
or only slightly swollen in the apices, without
internal pigment. Asci cylindrical or clavate,
50–68 µm long, 7–13 µm wide, I/KI + blue,
with indistinct or sometimes tapering ocular
chamber. Ascospores straight or slightly
curved, (40·0–)47·2–57·0–66·9(–82·0) µm
long, (2·50–)2·95–3·45–3·95(–4·50) µmwide
(n=66), with (6–)7–9–11(–15) septa (n=43).
Pycnidia immersed in the thallus, black,
50–100 µm diam. Conidia curved, non-sep-
tate, 14–17 × 1·0µm.
Etymology. The species is named with
reference to the thallus structure.
Habitat and distribution. The species is
known only from a single locality in Khabar-
ovskiy Krai, in a coniferous-broadleaf forest
with high humidity on a terrace above the river.
Comments.Bacidia areolata is very similar
to B. suffusa but differs by its smooth, cracked
to areolate thallus, lighter apothecia with less
developed and often inconspicuous white
pruina on the exciple edge, and shorter
spores. It can also be separated by the thinner
hymenium, which never exceeds 100 µm, but
this feature alone is not enough to confirm
the species. Bacidia suffusa from North
America differs by having larger pycnidia,
100–125 µm diam., with filiform curved,
non-septate conidia, 10–27 × 0·8µm.
Additional specimens examined. Only type material seen.
Bacidia elongata J. Gerasimova &
A. Beck sp. nov.
MycoBank No.: MB 821185
Similar to Bacidia fraxinea but differs in having a wide
zone of enlarged cell lumina along the edge of the
exciple.
Type: Russia, Khabarovskiy Krai, Khabarovskiy
Rayon, Bolshekhekhtsirskiy State Natural Reserve, 48°
25'N, 134°77'E, 160 m, coniferous-broadleaf forest, on a
terrace above the river, on bark of Acer mono, 5 Septem-
ber 2013, J. V. Gerasimova s. n. (M M-0182571—holo-
type; M M-0182572, LE L-13007—isotypes). GenBank
Accession no: MH048626
(Figs 3C & D, 4C & D)
Thallus poorly defined, thin to rather thick,
smooth to areolate, consisting of scattered or
contiguous, ±flattened or convex areoles;
or granular, consisting of ±globose, scattered
or cluster-forming granules; rarely cracked.
If the thallus forms a thick crust it is wrinkled
to warted, consisting of layered irregularly-
shaped warts; whitish when smooth, greyish,
light green, greyish green, dark grey-green,
with crystals in the upper cortex. Prothallus
inconspicuous, sometimes present in
between areoles, whitish. Photobiont chlor-
ococcoid green alga, 6–15 × 8–18 µm.
Apothecia (0·25–)0·40–0·55–0·70(–0·95)
mm, sessile, ±plane. Young apothecia occa-
sionally barrel-shaped, when mature ±plane,
to only slightly convex, rarely strongly convex
or irregularly shaped. Disc almost white or
pinkish in young apothecia, orange, orange-
brown to dark orange-brown and dark
purple-brown when mature. Margin con-
colourous or paler, light orange, rarely
brown; persistent, often with thick layer of
white pruina on the edge, especially in young
apothecia. Exciple 49·0–79·5–110·0µm wide,
without crystals, but sometimes with clusters
of crystals along the rim dissolving in N;
colourless to orange-brown, with 4 layers of
enlarged cells along the edge (1 layer of
terminal cells ±globose, up to 5·0×5·0–
7·0µm, other 3 layers ±cylindrical with
lumina that are 9–14(–20)×4·0–5·5µm).
Exciple rim laterally almost colourless, yel-
lowish, pale yellow-brown to orange-brown,
consisting of radially arranged hyphae. Mid-
dle exciple orange-brown to colourless, con-
sisting of periclinally arranged thin hyphae; K
+ yellow or brown, parts K+ purplish, N −.
Hymenium 62·5–92·0–110·0µm high, col-
ourless in lower part; upper part diffusely
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coloured, colourless to pale yellow, pale
orange, rarely brown-orange. Hypothecium
almost colourless to pale yellow, pale yellow-
brown to orange-brown, usually darker than
exciple below. Paraphyses simple, sometimes
fork-branched, some with unclear septa,
thin, 1·0–1·5–1·8µm wide in mid-hyme-
nium, ±clavate or only slightly swollen in the
apices, 2·0–2·5–3·0µm wide, without internal
pigment. Asci cylindrical, 43–72–95 µm long,
7–11–19 µm wide, I/KI+ blue, ocular cham-
ber inconspicuous. Ascospores acicular,
straight or slightly curved, (39–)51–59–68(–
80) µm long, (2·0–)2·5–3·0–3·5(–4·0) µm wide
(n=93), with (2–)5–7–12(–16) septa (n=71).
Pycnidia not seen.
Etymology. The species is named with
reference to the exciple structure, character-
ized by a wide zone of enlarged cell lumina
along the edge.
Habitat and distribution. Corticolous spe-
cies, occurring in mixed forests on the bark of
hardwoods. Known phorophytes: Acer mono,
Fraxinus mandshurica and Ulmus glabra.
Comments. There are some differences
between specimens of this species collected
in Khabarovskiy (holotype) and those from
Primorskiy Krai. The type specimen from
Khabarovskiy Krai differs partly in having
apothecia mainly without pruina, hypothe-
cium and exciple almost colourless or yel-
lowish K±intensifying, and only the rim is
coloured, brown-orange to orange-brown.
By contrast, the specimens from Primorskiy
Krai have a coloured hypothecium and exci-
ple, and apothecia primarily with a thick layer
of white pruina on the edge.
A pale specimen of B. elongata with a
mostly smooth thallus appears to be close to
B. fraxinea, but the wide zone of cells with
enlarged lumina along the edge of the exciple
clearly differentiates it from that species. In
fact, the zone of cells with enlarged lumina in
combination with its overall habit, places it
morphologically close to B. suffusa,B. mill-
egrana,B. campalea and related species. The
dark morph of B. elongata is similar to
B. suffusa but differs in lacking abundant
clusters of crystals in the exciple, a distinct
pigment in the exciple rim and having dif-
ferent cell size along the edge of the exciple.
Additional specimens examined.Russia: Primorskiy
Krai: Chuguyevskiy Rayon, Verkhneussuriyskiy Statsio-
nar, in the valley of the Sokolovka River, conifer-
broadleaf forest, on bark of Ulmus glabra, 1973, L. N.
Vasil’yeva s. n. (PIG 29682); Krasnoarmeiskiy Rayon,
forest close to the Mel’nichnoye settlement, birch forest
(Betula costata), on bark of Fraxinus mandshurica, 21 viii
2013, J. V. Gerasimova s. n. (M M-0182626, LE
L-13010); birch forest (Betula costata), on bark of Ulmus
glabra, 21 viii 2013, J. V. Gerasimova s. n.
(M M-0182627, LE L-13011); mixed forest with a pre-
dominance of Pinus koraiensis, with undergrowth of Acer
mono and Populus tremula, on bark of Acer mono, 22 viii
2013, J. V. Gerasimova s. n. (M M-0182628, LE
L-13012); mixed forest with a predominance of Pinus
koraiensis, with undergrowth of A. mono and Populus tre-
mula, on bark of F. mandshurica, 22 viii 2013, J. V.
Gerasimova s. n. (M M-0182625, LE L-13013); the
Sikhote-Alin’Nature Reserve, lowland forest, on bark of
A. mono, 30 vi 1977, I. F. Skirina s. n. (PIG 28762);
Partizanskiy Rayon, north-western slope of Mt. Lazovs-
kaya, 43°39'12·9''N, 133°35'48·0″E, 1132 m, spruce-fir
forest, on bark of Picea sp., 17 viii 2009, I. F. Skirina s. n.
(PIG 26560); Mt. Ol’khovaya, 540 m, coniferous-
broadleaf forest, on bark of Acer mandshuricum, 2010,
I. F. Skirina s. n. (PIG 29468); valley of Postyshevka
River, surroundings of the Krasnoarmeyskiy way station,
43°10'6·49''N, 133°00'9·35''E, 312 m, lowland forest, on
bark of Chosenia sp., 26 viii 2012, I. F. Skirina,F. V.
Skirin s. n. (PIG 32040).
Bacidia kurilensis J. Gerasimova,
A. Ezhkin & A. Beck sp. nov.
MycoBank No.: MB 821186
Similar to Bacidia laurocerasi but differs by the presence
of a green hue in the epihymenium and upper part of the
excipulum edge, as well as by a distinctly granular
thallus.
Type: Russia, Sakhalin Oblast, Kurile Islands, Kuna-
shir Island, at the foot of the Mendeleev Volcano, 44°
00'4·78''N, 145°42'26·85''E, 135 m, mixed conifer-
broadleaf forest, on bark of Salix udensis, 26 July 2013,
A. K. Ezhkin [B11/11.15] (M M-0182620—holotype;
SAK 276—isotype).
GenBank Accession no: MH048610
(Figs 3E & F, 4E & F)
Thallus poorly defined, thin to thick, partly
smooth, granular to granular isidiose; com-
posed of discrete or more often con-
tiguous, ±globose or extended, irregular
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granules, forming a loose assemblage, some-
times slightly flattened to subsquamulose;
light green-grey, grey-green, in the herbar-
ium becoming partly brownish; lacking crys-
tals in the upper cortex. Prothallus
epiphloeodal, often present between granules
or bordering the thallus, white or greyish.
Photobiont chlorococcoid green alga, 6–15 ×
8–18 μm and frequently with associated, free-
living cyanobacteria.
Apothecia (0·5–)0·7–0·9–1·2(–1·4) mm
diam., sessile, ±plane or very slightly convex,
becoming moderately convex when mature,
epruinose. Disc reddish brown, fuscous
brown to almost black, rarely mottled and
light brown in the middle. Margin con-
colorous with the disc, sometimes paler in the
lower part of young apothecia or reddish
brown. Exciple laterally 62·5–78·0–112·5µm
wide, without crystals. Rim dark brown in
upper part with a greenish hue, lower down
paler, brown to orange-brown; colourless in
inner part and under hypothecium; brown
pigment along the full length of the edge or
rim with a single layer of enlarged cells up to
6×5µm, without crystals, K+ intensifying.
Hymenium 80·0–95·0–107·5µm high, col-
ourless in lower part, without crystals; upper
part dark brown with a dirty green pigmen-
tation, K+ intensifying. Hypothecium pale
yellow, pale brown-yellow, almost colour-
less. Paraphyses simple, 1·5–2·0µm in mid-
hymenium, non-septate, slightly swollen at
apices 2·5–4·0µm, without internal pigment.
Asci clavate to cylindrical, 54–70 × 9–11 mm;
I/KI+ blue, ocular chamber inconspicuous.
Ascospores acicular, straight or slightly
curved, sometimes coiled in the ascus, (41–)
55–65–74(–88) µm long (n=85), (2·00–)
2·30–2·75–3·15(–4·00) µm wide, with (3–)5–
9–13(–17) septa (n=85).
Pycnidia not seen.
Etymology. The epithet ‘kurilensis’refers to
the group of islands where the species was
first collected.
Distribution and habitat. Known from
Kunashir Island at the foot of the Mendeleev
Volcano. It grows on the bark of Hydrangea
paniculata,Kalopanax septemlobus and Salix
udensis in a sparse conifer-broadleaf forest
with Abies sachalinensis and Picea jezoensis in a
small river valley. The habitat is associated
with high humidity and moderate insolation.
Comments.Bacidia kurilensis is closely
related to B. biatorina,B. heterochroa,
B. laurocerasi and B. salazarensis.Bacidia
biatorina has a similar thallus structure and
dark apothecia but differs in the shorter
spores (42–57 µm) and lack of green pig-
mentation in the epihymenium and edge of
the exciple. Bacidia salazarensis is character-
ized by spores having a lower length-width
ratio, a rimose thallus and a different dis-
tribution (the only Asian specimen of B. sal-
azarensis seen was from southern China).
Bacidia laurocerasi has a similar exciple
structure and long multiseptate spores but
differs by having a smooth to areolate thallus
and lacking the green pigmentation in the
exciple and epihymenium. Bacidia hetero-
chroa differs mainly by lacking the granular
thallus.
Additional specimens examined.Russia: Sakhalin:
Sakhalin Oblast, Kunashir Island, at the foot of the
Mendeleev Volcano, 44°00'4·78''N, 145°42'26·85''E,
135 m, mixed conifer-broadleaf forest, on bark of Kalo-
panax septemlobus, 2013, A. K. Ezhkin B7/11.15
(M M-0182621, LE, SAK 272); 44°00'4·78''N, 145°
42'26·85''E, 135 m, mixed conifer-broadleaf forest, on
bark of Hydrangea paniculata, 2013, A. K. Ezhkin B17/
11.15 (M M-0182622, LE, SAK 282).
Bacidia sachalinensis J. Gerasimova,
A. Ezhkin & A. Beck sp. nov.
MycoBank No.: MB 821187
Similar to Bacidia polychroa but differing in thallus
and exciple structure, and in its shorter spores with fewer
septa (40–58 × 2–3µm with 1–8 septa).
Type: Russia, Sakhalin, Sakhalin Oblast, Yuzhno-
Sakhalinsk, Rogatka River, 46°58'5·70''N, 142°
47'49·03''E, 163 m, floodplain forest, on bark of Populus
maximowiczii, 19 May 2014, A. K. Ezhkin [B8/12.14]
(M M-0182619—holotype; LE L-12961, SAK 145—
isotypes).
GenBank Accession no: MH048621
(Figs 3G & H, 4G & H)
Thallus poorly defined, thin to thick,
either discontinuous, smooth, indistinctly
areolate, consisting of scattered, discrete to
contiguous, ±flattened small areoles, or
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continuous, warted to wrinkled, cracked;
white, whitish green, pale grey to dirty grey,
grey-green. Prothallus sometimes present,
between areoles, white. Photobiont chlor-
ococcoid green alga, 6–15 × 8–18 μm.
Apothecia (0·30–)0·45–0·65–0·85(–1·30)
mm diam., ±plane when young, later
becoming convex, epruinose or rarely with
thin white pruina on the edge and the disc of
young to medium-aged apothecia. Disc pale
orange to intensely orange, brown-orange,
yellow-brown, rusty brown to dark orange-
brown and red-brown, rarely dark brown
when mature. Margin concolorous with the
disc or slightly darker, raised above disc in
young apothecia, later level with the disc,
and finallyexcludedinoldandconvexapo-
thecia. Exciple laterally 46·0–63·8–75·0µm
wide, without crystals. Rim pale brown,
yellow-brown, orange-brown, lower down
almost colourless, along the margin edge
with 2 layers of enlarged cells with lumina
that are 2·3–3·8–7·8×2·6–5·4–7·7µm(if±
globose then up to 8 × 8 µm), without crys-
tals; inner part (the same as rim) pale brown,
yellow-brown, orange-brown, sometimes
almost colourless below; K+ purplish.
Hymenium 70·0–78·8–92·5µm high, in lower
part colourless, in upper part indistinct and
diffusely coloured, pale yellow-brown, pale
orange-brown to orange-brown, sometimes
olive, yellowish, almost colourless. Hypo-
thecium pale brown-yellow to yellow-brown,
brown-orange, darker than exciple, K+
purplish. Paraphyses simple, thin, 1·60–
1·85–2·10 µm wide in mid-hymenium, ±
clavate or only slightly swollen in the apices,
1·8–2·4–3·5µm wide, without internal pig-
ment. Asci cylindrical or clavate, 37·0–54·5–
66·0×7·0–11·0–16·7µm wide, I/KI + blue,
with tapering ocular chamber. Ascospores
acicular, straight or slightly curved, some-
times coiled in ascus (36·7–)43·5–49·1–54·7
(–63·5) µmlong(2·00–)2·40–2·70–3·00(–-
4·25) µmwide(n=125), with (1–)3–5–7(–
8) septa.
Etymology. The epithet ‘sachalinensis’refers
to the locality where the species was collected.
Distribution and habitat. Known only from
a single locality on Sakhalin. It was collected
on the bark of mature trees, in an old flood-
plain poplar-willow forest with high under-
storey in a very humid habitat with a fair
amount of sunlight. Known phorophytes
include Populus maximowiczii and Ulmus
laciniata.
Comments.Bacidia sachalinensis has a very
variable thallus structure and apothecial
colour and this is also typical for North
American and European specimens of
B. polychroa. It is morphologically and ana-
tomically very similar to B. polychroa but dif-
fers in having 1–2 layers of cells with enlarged
lumina along the edge of the exciple, shorter
spores with fewer septa (40–58 × 2–3µm with
1–8 septa) and a usually smooth and poorly
defined thallus with light coloured apothecia.
North American specimens of B. polychroa
have longer and wider spores with more septa
(31–74 × 1·9–5·0µm with 2–15 septa (Ekman
1996)) while European specimens have
spores that are intermediate in size (33–
75 ×2·0–4·5µm with 3–16 septa (Foucard
2001; Llop 2007; Coppins & Aptroot 2009;
Wirth et al. 2013)). Bacidia diffracta differs
mainly by having a granular thallus.
Additional specimens examined.Russia: Sakhalin:
Sakhalin Oblast, Yuzhno-Sakhalinsk neighbourhood,
Rogatka River, 46°58'4·789''N, 142°48'18·88''E, 161 m,
floodplain forest, on bark of Populus maximowiczii, 2014,
A. K. Ezhkin B7/12.14 (M M-0182621, LE L-12960,
SAK 144); 46°58'5·707''N, 142°47'49·03''E, 163 m,
floodplain forest, on bark of Populus maximowiczii, 2014,
A. K. Ezhkin B9/12.14 (M M-0182623, LE L-12962,
SAK 146); 46°58'2·118''N, 142°46'16·75''E, 108 m,
floodplain forest, on bark of Populus maximowiczii, 2014,
A. K. Ezhkin B10/12.14 (LE L-12963, SAK 147); 46°
58'5·707''N, 142°47'49·03''E, 162 m, floodplain forest,
on bark of Ulmus laciniata, 2014, A. K. Ezhkin B11/12.14
(LE L-12964, SAK 148); 46°58'4·789''N, 142°
48'18·88''E, 161 m, floodplain forest, on bark of
Populus maximowiczii, 2014, A. K. Ezhkin B12/12.14
(M M-0182624, LE L-12965, SAK 149).
Bacidia schweinitzii (Fr.) A. Schneid.
Bacidia schweinitzii occurs in the temperate
forests of Canada around the Great Lakes
and the Maritimes but it also occurs in east-
ern Asia and the eastern parts of the USA as
far south as northern Florida (Ekman 1996;
Lendemer et al. 2016). The species is
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reported here from Russia for the first time. It
was collected in the temperate region of the
southern part of the Russian Far East, in
Primorskiy and Khabarovskiy Krai, and also
on Kunashir Island.
This species has been found on the bark
and trunks of a wide variety of conifer and
deciduous trees, often among or on top of the
branches colonized by mosses in dense
coniferous-broadleaf and spruce-fir forests.
At the Russian sites, the understorey often
has ferns, bryophytes and bamboo present.
Specimens were collected on several occa-
sions on the bark of fallen, well-decomposed
trees in shaded, very humid sites. Known
phorophytes in the Russian Far East include:
Abies nephrolepis,Acer ukurunduense,Betula
costata,B.ermanii,Fraxinus mandshurica,
Kalopanax septemlobus,Picea jezoensis,Picea
sp., Pinus sp., Populus tremula,Prunus cerasus,
Quercus mongolica,Quercus sp., Tilia amur-
ensis and Ulmus laciniata.
Specimens examined.Russia: Khabarovskiy Krai:
Khabarovskiy Rayon, Bolshekhekhtsirskiy Nature
Reserve, 48°25'N, 134°77'E, 157 m, coniferous-
broadleaf forest, on a terrace above the river, on bark of
Picea jezoensis, 5 ix 2013, J. V. Gerasimova s. n. (LE
L-13004); 48°22'N, 134°77'E, 865 m, spruce-fir forest
with Betula ermanii, on bark of Picea jezoensis, 3 ix 2013,
J. V. Gerasimova s. n. (M M-0182635, LE L-13003);
48°21'N, 134°79'E, 845 m, thick spruce forest, on bark
of Picea jezoensis, 4 ix 2013, J. V. Gerasimova s. n.
(M M-0182634, LE L-12999); 48°23'N, 134°77'E,
451 m, coniferous-broadleaf forest near the cordon, on
bark of Fraxinus mandshurica, 2 ix 2013, J. V. Gerasimova
s. n. (M M-0182580, M M-0182629, LE L-13001,
L-12998); 48°22'N, 134°77'E, 820 m, spruce forest with
Betula ermanii on the edge of drying zone, on bark of
Picea jezoensis, 3 ix 2013, J. V. Gerasimova s. n. (M M-
0182629, LE L-12998); ibid,J. V. Gerasimova s. n. (M
M-0182630, LE L-13000); 48°22'N, 134°77'E, 865 m,
spruce-fir forest with Betula ermanii, on bark of Abies
nephrolepis, 3 ix 2013, J. V. Gerasimova s. n. (M
M-0182631, LE, UPS L-721214); 48°23'N, 134°77'E,
451 m, coniferous-broadleaf forest near the cordon, on
bark of Betula costata, 2 ix 2013, J. V. Gerasimova s. n.
(LE L-13002); Kukanskiy Range, 50°55'N, 134°26'E,
715 m, spruce-larch green moss forest, on bark of Picea
sp., 23 viii 2012, I. A. Galanina,L. S. Yakovchenko s.
n. (VBGI). Primorskiy Krai: Chuguyevskiy Rayon,
Mt. Snezhnaya, south-west slope, old spruce-fir forest,
on top of moss twigs on bark of Picea sp., 5 viii 2003, I. F.
Skirina,F. V. Skirin s. n. (PIG 15673); Verkhneussur-
iyskiy Statsionar, in the valley of the Sokolovka River,
conifer-broadleaf forest, on bark of Acer ukurunduense,
1973, L. N. Vasil’yeva s. n. (PIG 13267);
Verkhneussuriyskiy Statsionar, in the valley of the
Sokolovka River, conifer-broadleaf forest, lowland for-
est, on bark of Prunus cerasus, 15 vii 1980, L. N. Vasi-
l’yeva s. n. (PIG 13268); Khasanskiy Rayon,
neighbourhood of Kravtsovka settlement, 42°38'N, 141°
44'E, oak (Quercus mongolica) forest, on bark of Quercus
mongolica, 6 v 2013, I. A. Galanina s. n. (VBGI); ibid., on
bark of Q. mongolica, 6 v 2013, I. A. Galanina s. n.
(VBGI); Ryazanovka River, oak forest, on bark of Quer-
cus sp., 1985, I. F. Skirina s. n. (PIG 5653); neighbour-
hood of Peschanyy Peninsula, oak forest, on bark of
Quercus sp., 17 viii 2008, I. F. Skirina s. n. (PIG 23811);
Krasnoarmeiskiy Rayon, western slope of the Sikhote-
Alin’, 46°13'N, 136°70'E, 1338 m, slope of the upper
reaches of Valincu River, on bark of Picea jezoensis, 25 viii
2013, J. V. Gerasimova s. n. (M M-0182579, LE, UPS
L-721217); 46°21'N, 136°66'E, 1010 m, thick spruce-fir
forest with fir and mosses in the understorey, on bark of
Abies nephrolepis, 29 viii 2013, J. V. Gerasimova s. n. (LE
L-12997); 46°15'N, 136°70'E, 1180 m, spruce-fir forest
with mosses in the understorey, on bark of Picea jezoensis,
26 viii 2013, J. V. Gerasimova s. n. (LE); 46°14'N, 136°
70'E, 1275 m, thick spruce-fir forest, the southern slope,
in the upper reaches of Valincu River, on bark of Picea
jezoensis, 26 viii 2013, J. V. Gerasimova s. n. (LE); ibid.,
on bark of Betula costata, 26 viii 2013, J. V. Gerasimova s.
n. (LE L-12996); western slopes of the Sikhote-Alin’,
upper reaches of the Bol’shaya Ussurka, the northern
slope, spruce-fir forest, on a fallen tree, 1981,
I. F. Skirina s. n. (PIG 6653); ibid., on bark of Acer
ukurunduense, 1981, I. F. Skirina s. n. (PIG 6600);
Sikhote-Alin’Nature Reserve, Sredniy Creek, birch
(Betula ermanii) forest, on bark of Betula ermanii, 1980,
I. F. Skirina s. n. (PIG 5578); Sikhote-Alin’Nature
Reserve, neighbourhood of Mt. Kolumbe, spruce-fir
forest, on bark of Abies sp. and Picea sp., 15 vii 1980, I. F.
Skirina s. n. (PIG 3437); Lazovskiy Rayon, neighbour-
hood of Valentin settlement, the Mt. Koldun rise,
coniferous-broadleaf forest, on bark of Quercus sp., 25
viii 2009, I. F. Skirina s. n. (PIG 24140); Partizanskiy
Rayon, Alekseyevskiy Range, Mt. Olkhovaya, the
southern slope, 700 m, conifer-broadleaf forest, on bark
of Quercus sp., 4 vii 2007, I. F. Skirina s. n. (PIG 21200);
Alekseyevskiy Range, Mt. Ol’khovaya, 540 m, lowland
forest near the river on bark of Populus tremula, 7 viii
2007, I. F. Skirina s. n. (PIG 23804); the spur of Mt.
Chantinza, 43°08'N, 132°58'E, 602 m, coniferous-
broadleaf forest, on bark of Tilia amurensis, 25 viii 2012,
I. F. Skirina,F. V. Skirin s. n. (PIG 32039); Pozharskiy
Rayon, spurs of Strel’nikov Range, 1 km from the out-
post, 263 m, oak forest, oak on the slope, on bark of
Quercus sp., 20 vii 2007, I. F. Skirina s. n. (PIG 21850);
Spasskiy Rayon, neighbourhood of Orlovka settlement,
45°20'46·5''N, 133°36'50·5''E, mixed forest, on bark of
Quercus sp., 26 vi 2009, I. F. Skirina,F. V. Skirin s. n.
(PIG 25692); Terneiskiy Rayon, Terneiskiy forestry,
neighbourhood of Tayozhnyy settlement, Lagernaya
River, 45°42'18·2''N, 136°17'40·3''E, 717 m, pine-spruce
forest on the west-south-west slope, on bark of Pinus sp.,
10 vii 2011, I. A. Galanina s. n. (VBGI); neighbourhood
of Tayozhnyy settlement, Mrac hnyy Creek, 45°44'38·9''N,
136°09ʹ29·3''E, 745 m, pine-spruce forest, on bark of Picea
622 THE LICHENOLOGIST Vol. 50
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jezoensis, 9 vii 2011, I. A. Galanina s. n. (VBGI); neigh-
bourhood of Tayozhnyy settlement, 45°41'07·4''N, 136°
10'26·5''E, 713m, pine-spruce forest on a gentle slope, on
the top of moss twigs on bark of Picea sp., 8 vii 2011, I. A.
Galanina s. n. (VBGI); neighbourhood of Tayozhnyy
settlement, Mrachnyy Creek, 45°44'13·6''N, 136°
09'31·9''E, 745m, pine-spruce forest, on bark of Picea
jezoensis, 2011, I. A. Galanina s. n. (VBGI); ibid., on bark
of Picea jezoensis, 2011, I. A. Galanina s. n. (VBGI). The
Jewish Autonomous Oblast: Bastak Nature Reserve,
“Dubovaya Sopka”, near the cordon, oak forest, on bark
of Quercus sp., 17 x 2005, I. F. Skirina s. n. (PIG 18012).
Sakhalin: Sakhalin Oblast, Kunashir Island, at the foot of
the Mendeleev Volcano, 43°59'37·07''N, 145°46'50·62''E,
105 m, old spruce-fir forest with Abies sachalinensis,Picea
glehnii and P. jezoensis, on bark of fallen strongly decom-
posing tree, 2014, A. K. Ezhkin B20/11.15 (SAK 285);
neighbourhood of Lagunnoye Lake, 44°02'50·2''N, 145°
46'01·6''E, 79m, mixed conifer-broadleaf forest, on bark
of Kalopanax septemlobus, 2015, A. K. Ezhkin B10/11.15
(SAK 275); neighbourhood of Lagunnoye Lake, 44°
02'50·2''N, 145°46'01·6''E, 79m, mixed conifer-broadleaf
forest, on bark of Ulmus laciniata, 2015, A. K. Ezhkin B21/
11.15 (SAK 286).
We thank S. Ekman (Uppsala, Sweden) for many helpful
comments on an earlier draft of this manuscript and
advice on characters of the new Bacidia species. We are
grateful to David Richardson (Halifax, Canada) and the
editors of the Lichenologist for their corrections to the
English text. The curators of LE, MIN, UPS, US, VBGI,
Pacific Institute of Geography, as well as I. A. Galanina
and I. F. Skirina (both Vladivostok, Russia), provided
herbarium material used in this study. Our thanks to the
late J. Heinrichs (Munich, Germany) for access to pho-
tographic equipment and J. Bechteler (Munich, Ger-
many) for assistance at the early stage of laboratory work.
The field work was supported by the World Wildlife
Fund (WWF352/RU009622/GLM) within the project
“Study of the indicator species of lichens and fungi in the
dark conifer forests”and the Russian Foundation for
Basic Research (no. 14-04-01411). Molecular work was
supported by a grant from the Bayerisches Staatsminis-
terium für Bildung und Kultus, Wissenschaft und Kunst
in the “Barcoding Fauna Bavarica”framework, and a
BAYHOST fellowship (MB-2017-1/35) to the last
author.
SUPPLEMENTARY MATERIAL
For supplementary material accompanying this paper
visit https://doi.org/10.1017/S0024282918000397
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