The taxonomy of the Caloplaca citrina group (Teloschistaceae) in the Black Sea region; With contributions to the cryptic species concept in lichenology

Abstract

A new taxonomy of the Caloplaca citrina group in the Black Sea region is presented. It is based on the nrDNA ITS molecular data, chemistry (anthraquinone contents) and 20 morphological characters. Six species previously known in the region are accepted: Caloplaca arcis , C. calcitrapa , C. dichroa , C. flavocitrina , C. geleverjae , C. limonia . Five new species are described: Caloplaca arcisproxima , C. austrocitrina , C. communis , C. confusa and C. nigromarina . Seven further species, Caloplaca britannica , C. citrina , C. marina , C. maritima , C. microthallina , C. ora and C. phlogina are also treated briefly. Some maritime species known from the Atlantic coast of Europe are absent from the region, and, surprisingly, Caloplaca citrina s. str. could not be confirmed from the study area. A key to the species present in the region is provided, although morphological characters are of very limited value in this group. The variability and taxonomic importance of particular features are discussed. No significant differences in secondary chemistry were observed among the species.
Many examples of convergence and some semi-cryptic species were revealed by molecular data. The term ‘semi-cryptic species’ is introduced here into lichenology for those species which cannot be clearly diagnosed by their morphology, but which are determined by other characters, mainly by their ecology and distribution. We propose to describe formally such species, in spite of difficulties with subsequent morphological identification.

Full text

The taxonomy of the Caloplaca citrina group
(Teloschistaceae) in the Black Sea region; with contributions
to the cryptic species concept in lichenology
Jan VONDRÁK, Pavel R
ˇÍHA, Ulf ARUP and Ulrik SØCHTING
Abstract: A new taxonomy of the Caloplaca citrina group in the Black Sea region is presented. It is
based on the nrDNA ITS molecular data, chemistry (anthraquinone contents) and 20 morphological
characters. Six species previously known in the region are accepted: Caloplaca arcis,C. calcitrapa,
C. dichroa,C. flavocitrina,C. geleverjae,C. limonia. Five new species are described: Caloplaca
arcisproxima,C. austrocitrina,C. communis,C. confusa and C. nigromarina. Seven further species,
Caloplaca britannica,C. citrina,C. marina,C. maritima,C. microthallina,C. ora and C. phlogina are also
treated briefly. Some maritime species known from the Atlantic coast of Europe are absent from the
region, and, surprisingly, Caloplaca citrina s. str. could not be confirmed from the study area. A key to
the species present in the region is provided, although morphological characters are of very limited
value in this group. The variability and taxonomic importance of particular features are discussed. No
significant differences in secondary chemistry were observed among the species.
Many examples of convergence and some semi-cryptic species were revealed by molecular data. The
term ‘semi-cryptic species’ is introduced here into lichenology for those species which cannot be clearly
diagnosed by their morphology, but which are determined by other characters, mainly by their ecology
and distribution. We propose to describe formally such species, in spite of difficulties with subsequent
morphological identification.
Key words: alveolate cortex, Caloplaca citrina clade, lichens, nrDNA ITS, semi-cryptic species,
Teloschistales
Introduction
A number of lichenological papers, dealing
with taxonomy or biodiversity, have included
lichen species inhabiting seashore habitats
but only some of these have dealt with par-
ticular taxonomic groups occurring specifi-
cally on coastal cliffs, e.g. Ramalina in Sheard
(1978), Roccella in Tehler et al. (2004), Ver-
rucaria in for example Brodo & Santesson
(1997), Harada (2004) or McCarthy (1991),
and Xanthoria in Lindblom & Ekman
(2005). Marine and maritime lichens are
well-known in some regions, for instance in
the British Isles, where some ecological and
generalized studies have been made (e.g.
Fletcher 1973a,b, 1975a,b). The genus Calo-
placa has been intensively investigated on
North American shores (Arup 1992a,b,
1993a,b, 1994, 1995a,b, 1997b) and in
Europe (e.g. Tavares 1956; Laundon 1992;
Roux & Navarro-Rosinés 1992; Navarro-
Rosinés & Roux 1993, 1995; Arup 1997a).
In the East Mediterranean and the Black Sea
region, some papers, mainly contributions on
lichen biodiversity, partially cover maritime
species, including Caloplaca (e.g. Szatala
1943a,b;Veˇzda 1975; Roux & Navarro-
Rosinés 1992; Güvenc & Öztürk 1999;
Sipman & Raus 1999, 2002; Yazici 1999;
Khodosovtsev 2001, 2002, 2003; Redchenko
J. Vondrák: Department of Botany, Faculty of Science,
University of South Bohemia, Branišovská 31, C
ˇeské
Budeˇjovice, CZ-370 05, Czech Republic. Email:
j.vondrak@seznam.cz
P. R
ˇı´ha: Department of Zoology, Faculty of Science,
University of South Bohemia, Branišovská 31, C
ˇeské
Budeˇjovice, CZ-370 05, Czech Republic.
U. Arup: Botanical Museum, Lund Univeristy, Östra
Vallgatan 18, SE-223 61 Lund, Sweden.
U. Søchting: Section for Ecology and Evolution,
Department of Biology, University of Copenhagen, Ø.
Farimagsgade 2D, DK-1353 Copenhagen, Denmark.
The Lichenologist 41(6): 571–604 (2009) © British Lichen Society, 2009
doi:10.1017/S0024282909008317

2002; Khodosovtsev et al. 2003, 2007; John &
Breuss 2004; Sipman et al. 2005; Vondrák &
Slavı´ková-Bayerová 2006; Vondrák et al.
2008a;), but otherwise knowledge of mari-
time species of this area remains sparse.
While working in the Black Sea region, one
of us (JV) found that current W European
literature for the identification of lichens
(species in the genera Caloplaca,Candelari-
ella,Catillaria,Lecania and Toninia) was
inadequate. This was particularly true for
the genus Caloplaca, which appeared to be
extremely species-rich in the area. Currently,
we estimate that more than 50 species occur
in coastal habitats around the Black Sea.
This huge, but largely unknown diversity, led
us to initiate a complex project on the bio-
diversity of the genus Caloplaca in the Black
Sea region and the present study focuses on
the taxonomy of the Caloplaca citrina group in
the area. Arup (2006a) has already clarified
the taxonomy of this group in Scandinavia
and his results are utilized and developed
further here. This group seems to be rather
species rich and according to phylogenetic
studies it is closely related to the C. saxicola
group, various clades of Xanthoria species
and the C. holocarpa group (Arup & Grube
1999; Gaya et al. 2008).This paper describes
the species diversity in the C. citrina group.
The taxonomy proposed in this study is
based on molecular data using the single
locus, ITS nrDNA. This locus is usually well
suited for taxonomic studies at the species
level, and a number of published papers
using only this region has given meaningful
results in the genus Caloplaca (Tretiach et al.
2003; Arup 2006a, 2009; Arup et al. 2007;
Søchting & Figueras 2007; Muggia et al.
2008; Vondrák et al. 2008b,c). The only
study in Teloschistaceae so far using a multi-
locus approach is one dealing with taxonomy
at higher than species level (Søchting &
Lutzoni 2003). Our investigations have some
general implications exceeding the frame-
work of a regional taxonomic study since
the molecular results have revealed several
distinct groups, which are often morphologi-
cally hardly distinguishable. This phenom-
enon, termed semi-cryptic speciation, is
discussed at the end of the paper.
Material and Methods
Material from the Atlantic coast of Europe used for
comparison (mainly of Caloplaca britannica,C. littorea,
C. marina,C. maritima,andC. microthallina) was ob-
tained on loan from BM, CBFS, LD, and hb. A. Aptroot
(ABL). Material from Central Europe, the Black Sea
region and the Mediterranean was obtained on loan
from B, BP, CBFS, GZU, KHER, PRM, SAV and W.
The main part of the material from the Black Sea region
was collected by the first author during field excursions
in July 2004 (Bulgaria), July and November 2005
(Romania, Bulgaria, European part of Turkey), June
2006 (Ukraine: Crimea), and April, May 2007
(Romania, Bulgaria, Turkey, Georgia, Russia, Ukraine).
Vouchers are deposited in CBFS (the numbers follow-
ing this abbreviation are herbarium accession numbers).
Citations of specimens examined are abbreviated and
for common species only selected specimens are pre-
sented. Data on additional samples and full sample
information are available on the web page: http://
botanika.bf.jcu.cz/lichenology/data.php.
Morphology and anatomy
A total of 20 characters was measured and used in the
detailed characterization of the Caloplaca citrina group,
but only 11 traits were important for species separation:
width of areoles, thickness of thallus and cortex/alveolate
cortex, size of vegetative diaspores, size of apothecia,
exciple width, hymenium height, size of ascospores,
width of spore septa, width of paraphyses tips and size of
conidia.
Sections for morphological examination were cut by
hand and observed in water, but paraphysis tips and
cortical tissues were observed after pretreatment with
KOH. Measurements were made to an accuracy of
0·5 µm for cells (e.g. ascospores, conidia and para-
physes), 1 µm for thickness of the cortex and 10 µm for
larger structures (e.g. hymenium thickness and exciple
width). All measurements of cells included their walls.
Measurements are given as (min.–) x± SD (–max.),
where x= mean value and SD = standard deviation. The
total numbers of measurements (n) are given in paren-
theses. At least five measurements were taken from all
specimens available; in the morphologically indistin-
guishable species (Caloplaca confusa,C. nigromarina),
only specimens confirmed by molecular data were used
for measurements.
Morphological terminology follows Ryan et al. (2002)
and Bungartz (2002). The term ‘alveolate cortex’ is
proposed here for a hyaline tissue formed by living
fungal cells among dead algal cells or in gaps left by dead
algal cells. It is similar to a phenocortex (sensu Ryan et al.
2002: 23), but differs in fungal cells, which are living,
and not dead as in a phenocortex. Being situated be-
tween algal and epinecral layers, it replaces a true cortex
in some species.
DNA extraction and amplification.
Direct PCR was used for DNA extraction and PCR-
amplification of the nuclear ITS regions including the
572 THE LICHENOLOGIST Vol. 41

T 1.Voucher specimens and GenBank accession numbers of the ITS sequences used in the phylogenetic analysis. Accession
numbers in bold represent new sequences produced during this study
Species & Herbarium
Accession No.
Source GenBank
Accession No.
Caloplaca arcis Austria. (Arup 2006a) DQ173213
C. arcis Sweden. (Arup 2006a) DQ173214
C. arcis Great Britain, England. (Arup 2006a) DQ173215
C. arcis W1990-00525 Italy, Sardinia (coll. W. Brunnbauer 1986) EU563454
C. arcis CBFS JV3036 Bulgaria, Black Sea coast (coll. Vondrák 2005) EU563395
C. arcis CBFS JV4985 The Netherlands, Noordoostpolder (coll. A. Aptroot
59871, 2004)
EU563453
C. arcis CBFS JV4986 The Netherlands, Gelderland (coll. A. Aptroot 59582,
2003)
EU563452
C. arcis CBFS JV5426 Turkey, Black Sea coast (coll. Vondrák 2007) EU563424
C. arcis CBFS JV6093 Bulgaria, Black Sea coast (coll. Vondrák 2007) EU563438
C. arcisproxima CBFS JV4125 Greece, South Crete (coll. Vondrák 2005) EU563413
C. arcisproxima CBFS JV5473 Ukraine, Black Sea coast (coll. Vondrák 2007) EU563425
C. austrocitrina CBFS JV991 Czech Republic, C
ˇeské Budeˇjovice (coll. Vondrák 2003) EU563450
C. austrocitrina CBFS JV3436 Romania, Black Sea coast (coll. Vondrák 2005) EU563406
C. austrocitrina CBFS JV4195 Greece, North Crete (coll. Vondrák 2005) EU563416
C. austrocitrina CBFS JV4631 Bulgaria, Black Sea coast (coll. Vondrák 2005) EU563417
C. austrocitrina CBFS JV5236 Ukraine, Black Sea coast (coll. Vondrák 2006) EU563419
C. austrocitrina CBFS JV5285 Ukraine, Black Sea coast (coll. Vondrák 2006) EU563420
C. austrocitrina CBFS JV5474 Russia, Black Sea coast (coll. Vondrák 2007) EU563426
C. austrocitrina CBFS JV5476 Ukraine, Black Sea coast (coll. Vondrák 2007) EU563427
C. austrocitrina CBFS JV6097 Russia, Black Sea coast (coll. Vondrák 2007) EU563441
C. calcitrapa France, Languedoc-Roussellon, isotype (Arup 2006a) DQ173227
C. calcitrapa CBFS JV3408 Turkey, Black Sea coast (coll. Vondrák 2005) EU563401
C. calcitrapa CBFS JV5486 Bulgaria, Black Sea coast (coll. Vondrák 2007) EU563431
C. calcitrapa CBFS JV6100 Turkey, Marmara Sea coast (coll. Vondrák 2007) EU563444
C. citrina Sweden. (Arup 2006a) DQ173222
C. citrina Sweden. (Arup 2006a) DQ173223
C. citrina Sweden. (Arup 2006a) DQ173224
C. citrina Sweden. (Arup 2006a) DQ173225
C. citrina Sweden. (Arup 2006a) DQ173226
C. citrina CBFS JV1138 Czech Republic, Andeˇlská Hora (coll. Vondrák 2003) EU563387
C. communis CBFS JV3471 Turkey, Black Sea coast (coll. Vondrák 2005) EU563409
C. communis CBFS JV3042 Turkey, Black Sea coast (coll. Vondrák 2005) EU563397
C. communis CBFS JV3037 Turkey, Black Sea coast (coll. Vondrák 2005) EU563396
C. communis CBFS JV3367 Turkey, Black Sea coast (coll. Vondrák 2005) EU563399
C. communis CBFS JV3763 Greece, South Crete (coll. Vondrák 2005) EU563410
C. communis CBFS JV3803 Greece, North Crete (coll. Vondrák 2005) EU563411
C. communis CBFS JV5481 Turkey, Gallipoli Peninsula (coll. Vondrák 2007) EU563429
C. communis CBFS JV6092 Turkey, Gallipoli Peninsula (coll. Vondrák 2007) EU563437
C. communis CBFS JV6104 Russia, Taman Peninsula, Sea of Azov coast (coll.
Vondrák 2007)
EU563446
C. communis CBFS JV6113 Bulgaria, Black Sea coast (coll. Vondrák 2007) EU563447
C. communis CBFS JV6119 Turkey, Black Sea coast (coll. Vondrák 2007) EU563448
C. confusa CBFS JV3435 Bulgaria, Black Sea coast (coll. Vondrák 2005) EU563405
C. confusa CBFS JV6206 Bulgaria, Black Sea coast (coll. Vondrák 2007) EU563449
C. confusa herb. F. Berger Azores, Sao Jorje (coll. F. Berger 1992) EU563468
C. confusa GZU Haf31862 France, Corsica (coll. J. Hafellner 1993) EU563457
C. confusa GZU Italy, Sicily (coll. J. Poelt 1992) EU563455
C. dichroa Sweden. (Arup 2006a) DQ173228
C. dichroa Sweden. (Arup 2006a) DQ173229
C. dichroa Sweden. (Arup 2006a) DQ173230
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 573

T 1.Continued
Species & Herbarium
Accession No. Source GenBank
Accession No.
C. dichroa Sweden. (Arup 2006a) DQ173231
C. dichroa Sweden. (Arup 2006a) DQ173232
C. dichroa CBFS JV5324 Romania, Black Sea coast (coll. Vondrák 2007) EU563421
C. dichroa CBFS JV5477 Ukraine, Karadag (coll. Vondrák 2007) EU563428
C. dichroa CBFS JV4155 Great Britain, England (coll. Vondrák 2006) EU563415
C. dichroa GZU Haf43519 Austria, Tirolia, alt. c. 1000 m (coll. J. Hafellner 1997) EU563458
C. flavocitrina Sweden. (Arup 2006a) DQ173216
C. flavocitrina Sweden. (Arup 2006a) DQ173217
C. flavocitrina Sweden. (Arup 2006a) DQ173218
C. flavocitrina Sweden. (Arup 2006a) DQ173219
C. flavocitrina Sweden. (Arup 2006a) DQ173220
C. flavocitrina Austria. (Arup 2006a) DQ173221
C. flavocitrina CBFS JV1495 Czech Republic, Husinec (coll. Vondrák 2002) EU563388
C. flavocitrina CBFS JV2106 Bulgaria, Black Sea coast, epiphytic (coll. Vondrák 2004) EU563390
C. flavocitrina CBFS JV2536 Czech Republic, Kalubice (coll. Vondrák 2005) EU563392
C. flavocitrina CBFS JV3425 Bulgaria, Black Sea coast, concrete (coll. Vondrák 2005) EU563404
C. flavocitrina CBFS JV6089 Russia, Black Sea coast (coll. Vondrák 2007) EU563434
C. flavocitrina CBFS JV6090 Turkey, Black Sea coast (coll. Vondrák 2007) EU563435
C. flavocitrina CBFS JV6094 Georgia, Shuakhevi (coll. Vondrák 2007) EU563439
C. flavocitrina CBFS JV6095 Russia, Black Sea coast (coll. Vondrák 2007) EU563440
C. flavocitrina CBFS JV6098 Georgia, Black Sea coast (coll. Vondrák 2007) EU563442
C. flavocitrina GZU Haf31888 Italy, Livorno, coastal rock (coll. J. Hafellner 1993) EU563456
C. flavocitrina GZU Wetm76242 Hawaii, alt. c. 1700 m (coll. C. Wetmore 1996) EU563471
C. geleverjae CBFS JV5415 Ukraine, Black Sea coast, topotype (coll. Vondrák 2007) EU563423
C. havaasii BG Norway, topotype (Arup 2006b) DQ647649
C. limonia GZU Italy, Marettimo, paratype (coll. Poelt 1991) EU563467
C. limonia CBFS JV2515 Czech Republic, Pı´sek (coll. Vondrák 2005) EU563391
C. limonia CBFS JV3388 Bulgaria, Black Sea coast (coll. Vondrák 2005) EU563400
C. limonia CBFS JV3410 Turkey, Black Sea coast (coll. Vondrák 2005) EU563402
C. limonia CBFS JV3413 Turkey, Black Sea coast (coll. Vondrák 2005) EU563403
C. limonia CBFS JV3438 Bulgaria, Black Sea coast (coll. Vondrák 2005) EU563407
C. limonia CBFS JV3465 Bulgaria, Black Sea coast (coll. Vondrák 2005) EU563408
C. limonia CBFS JV5352 Romania, Black Sea coast (coll. Vondrák 2007) EU563422
C. limonia CBFS JV6101 Turkey, Black Sea coast (coll. Vondrák 2007) EU563445
C. marina Great Britain, England (Arup & Grube 1999) AF353946
C. marina North America, western coast (Arup & Grube 1999) AF353947
C. marina BM 730882 Great Britain, Wales (2004) EU563461
C. marina C US7520 Iceland, S-Múlasýla (coll. U. Søchting 1997) EU563470
C. maritima Great Britain, Wales (Arup & Grube 1999) AF353948
C. maritima CBFS JV4943 Great Britain, Wales (coll. U. Arup L92256, 1992) EU563462
C. maritima CBFS JV4987 The Netherlands, Walcheren (coll. A. Aptroot 59408,
2003)
EU563451
C. microthallina C US7480 Sweden, Halland (coll. U. Søchting 1997) EU563469
C. microthallina CBFS JV4939 Great Britain, England (coll. U. Arup L92274, 1992) EU563464
C. microthallina CBFS JV4940 Great Britain, England (coll. U. Arup L92291, 1992) EU563463
C. microthallina CBFS JV4941 Great Britain, Wales (coll. U. Arup L92307, 1992) EU563465
C. nigromarina GZU Lich.Anat.
Exsic. 80
Turkey, Black Sea coast (coll. V. John & E. Sauer 1992) EU563459
C. nigromarina CBFS JV3035a Turkey, Black Sea coast (coll. Vondrák 2005) EU563393
C. nigromarina CBFS JV3035b Turkey, Black Sea coast (coll. Vondrák 2005) EU563394
C. nigromarina CBFS JV3354 Turkey, Black Sea coast (coll. Vondrák 2005) EU563398
C. nigromarina CBFS JV5482 Turkey, Gallipoli Peninsula (coll. Vondrák 2007) EU563430
C. nigromarina CBFS JV4983 Bulgaria, Black Sea coast (coll. Vondrák 2004) EU563418
574 THE LICHENOLOGIST Vol. 41

5.8S region of the nuclear ribosomal DNA gene (abbre-
viated as ITS) following Arup (2006a). Primers for
amplification were ITS1F (Gardes & Bruns 1993) and
ITS4 (White et al. 1990). PCR cycling parameters fol-
lowed Ekman (2001).
Phylogenetic analyses
The ITS sequences used in the phylogenetic analysis
are listed in Table 1. Caloplaca holocarpa was used as the
outgroup. The alignment was conducted using MAFFT
6 (on-line version in the Q-INS-i mode; see Katoh et al.
2002) and manually trimmed in BioEdit 7.0 (Hall 1999)
to eliminate the unalignable ends.
Bayesian phylogenetic analysis was carried out with
a Metropolis-coupled Markov chain Monte Carlo
(MCMCMC) algorithm (Altekar et al. 2004) imple-
mented in MrBayes 3.0 (Ronquist and Huelsenbeck
2003). Prior to the analysis, Mr Modeltest 2.2 (Nylander
et al. 2004) was employed to assign the best-fit likelihood
settings according to the hLRT (likelihood ratio test).
The general time reversible model with some invariable
sites (GTR+I+) was determined as the best-fitting. A
flat Dirichlet prior distribution with all values set to 1·0
was used to model the prior probability densities of the
substitution rates as well as the stationary nucleotide
frequencies. In order to assess the stability of the
MCMC process, we monitored the standard deviation
of split frequencies of two simultaneous independent
runs, each including six parallel chains (one ‘cold’ and
five incrementally heated by a temperature of 0·3). Every
100th count of the total of 5 000 000 generations for
each run was sampled and the first quarter of samples
was discarded as burn-in. The resulting 75 000 trees
were used for reconstruction of a 50% majority-rule
consensus tree.
The heuristic parsimony search was conducted in
PAUP 4.0b10 (Swofford 2002), assessing the credibility
of branches via non-parametric bootstrapping. In order
to reduce the computational demands, a two-level
analysis was chosen: the treespace was examined during
the first run using the nChuck=3chuckScore=1 op-
tion for 100 replicates of the random sequence addition.
The second search was performed on the stored trees
and filled up the treespace thanks to the option
nChuck=0chuckScore=0. The maximum number of
trees in memory was restricted to 250 000 using the
maxTrees option. Bootstrap analysis encompassed
1000 resamplings and proceeded similar to the heuristic
parsimony search with the exception of nChuck setting
(=10) and number of RAS (=10) during the first run of
analysis. The TBR algorithm was employed, all charac-
ters were equally weighted and the gaps were treated as
missing data. The steepest descent option was not in
effect and the analysis ran under the mulTrees option.
Chemistry
The acetone extracts of anthraquinones were sub-
jected to high-performance liquid chromatography
(HPLC) analysis. Reverse phase column (C18, 5µm,
Lichrocart 250-4) was eluted with MeOH / 30%MeOH
+ 1%H
3
PO
4
for 77 min and the absorbances at 270 nm
were recorded (for details see Søchting 1997). The
compounds were determined on the basis of their reten-
tion times and absorption spectra; samples examined are
listed in Table 2.
Results and Discussion
Phylogeny
We confined the ‘Caloplaca citrina clade’ to
a group of morphologically variable species,
to which we found C. holocarpa as closest
outgroup (Fig. 1). According to Gaya et al.
(2008), Caloplaca granulosa is a closer relative
to the ‘Caloplaca citrina clade’, but it is prob-
ably too close to the ingroup or may even
belong within it. In fact, our circumscription
of the ‘Caloplaca citrina clade’ was made ar-
bitrarily to cover the majority of taxa with the
‘Caloplaca citrina morphology’ (see Morpho-
logy below) in the Black Sea region.
The resulting dataset contained 105 termi-
nals assigned to 20 species, of which 17 are
named here. The alignment had 674 posi-
tions, of which 233 were variable and 112
were parsimony informative. The first run of
T 1.Continued
Species & Herbarium
Accession No. Source GenBank
Accession No.
C. nigromarina CBFS JV6091 Georgia, Black Sea coast (coll. Vondrák 2007) EU563436
Caloplaca cf. ora CBFS JV3836 Greece, North Crete (coll. Vondrák 2005) EU563412
C. phlogina CBFS JV3437 Romania, Black Sea coast (coll. Vondrák 2005) EU563460
Caloplaca sp. CBFS JV4146 Greece, South Crete (coll. Vondrák 2005) EU563414
Caloplaca sp. CBFS JV6086 Turkey, Black Sea coast (coll. Vondrák 2007) EU563433
Caloplaca sp. CBFS JV2051 Bulgaria, Lyubimets (coll. Vondrák 2004) EU563389
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 575

T 2.Anthraquinone contents (% concentrations) of selected species in the Caloplaca citrina clade
teloschistin falacinal parietinic acid emodin parietin
C. arcis
CBFS JV3036 (AP) 0·6 2·3 1·1 1·0 93·9
CBFS JV3036 (TH) 1·0 8·1 1·0 0·7 89·3
CBFS JV5426 (AP) 1·0 7·5 1·4 0·4 88·8
C. austrocitrina
CBFS JV991 (TH) 1·4 5·4 0·0 1·0 92·2
CBFS JV4195 (AP) 3·3 16·8 2·1 1·3 76·5
CBFS JV4195 (TH) 3·4 26·7 1·3 1·5 67·1
CBFS JV5474 (AP) 2·3 14·1 1·6 0·8 80·1
C. calcitrapa
CBFS JV3408 (AP) 0·6 2·0 0·7 0·8 96·0
C. citrina s.str.
CBFS JV1138 (AP) 0·6 1·6 0·6 0·7 96·5
C. communis
CBFS JV3367 (AP) 0·7 1·6 0·9 0·8 96·0
CBFS JV3367 (TH) 1·4 2·1 0·0 2·1 94·4
CBFS JV3467 (AP) 2·0 4·6 1·1 0·7 91·6
CBFS JV3763 (AP) 2·2 6·2 1·9 0·8 88·6
CBFS JV3803 (AP) 2·3 4·1 1·3 0·6 91·5
CBFS JV4114 (AP) 4·6 6·7 1·7 0·5 86·5
C. confusa
CBFS JV3435 (TH) 1·2 3·8 1·0 1·0 93·0
C. dichroa
CBFS JV5337 (AP) 1·2 2·5 1·0 1·1 94·3
CBFS JV5337 (TH) 0·6 4·1 0·7 0·6 93·9
C. flavocitrina
CBFS JV3425 (AP) 5·3 12·1 1·4 0·7 80·4
CBFS JV3425 (TH) 1·5 9·6 0·0 0·0 88·8
C geleverjae
CBFS JV5415 (AP) 1·1 2·2 0·6 1·4 94·7
C. limonia
CBFS JV2515 (AP) 0·4 1·2 0·5 0·5 96·3
CBFS JV2515 (TH) 0·4 1·0 0·0 0·7 95·0
CBFS JV3388 (AP) 0·5 2·4 0·6 0·6 95·9
CBFS JV3388 (TH) 0·9 2·9 1·4 1·9 92·9
CBFS JV3438 (AP) 0·9 3·0 1·7 0·9 93·4
CBFS JV3465 (AP) 0·5 1·8 0·8 0·8 96·0
C. nigromarina
CBFS JV3035 (TH) 1·2 9·4 0·9 0·7 87·8
CBFS JV3354 (TH) 1·4 2·5 0·3 0·7 95·1
CBFS JV3399 (TH) 1·6 13·6 0·5 0·5 83·7
CBFS JV3478 (TH) 1·4 6·2 0·7 0·8 90·9
GZU, Lich. Anatol. Exs. 80 (AP) 1·4 5·9 2·0 1·1 89·7
GZU, Lich. Anatol. Exs. 80 (TH) 2·2 12·8 0·0 0· 85·0
C. ora
GZU, isotype (AP) 0·7 1·9 0·8 0·6 96·0
GZU, isotype (TH) 0·0 0·0 0·0 0·9 99·1
GZU, A. Veˇzda: Lich. Rar. Esc. 34 (AP) 0·4 1·3 0·9 0·4 97·0
GZU, A. Veˇzda: Lich. Rar. Exs. 34 (TH) 0·4 1·3 0·7 0·4 97·1
C. phlogina
CBFS JV3437 (AP) 0·6 2·2 1·2 0·7 95·3
CBFS JV3437 (TH) 0·6 2·2 1·2 0·7 95·3
576 THE LICHENOLOGIST Vol. 41

parsimony analysis yielded 67 islands of
equally parsimonious trees with the length
of 417 steps (due to the nchuck setting, each
island contains 3 trees and may in fact repre-
sent a fraction of a larger island). The con-
sistency index (CI) of the trees was 0·695 and
the retention index (RI) 0·640. The boot-
strap tree showed 40 supported internodes
(BS > 50%). The phylogenetic tree, based on
the Bayesian inference analysis is presented
here (Fig. 1) along with the posterior
probabilities (PP) for the nodes. Since the
parsimony bootstrap search yielded a tree
topologically congruent with Fig. 1, yet less
resolved (there were 52 internodes supported
by the Bayesian inference), we are adopting
only bootstrap values (BS) from this analysis.
Two groups, recognized by the parsimony
bootstrapping, but collapsed in the Bayesian
inference (clade with C. arcisproxima and a
grouping of three C. austrocitrina sequences),
are also depicted in Fig. 1, using additional
brackets. Molecular data revealed eleven,
partly semi-cryptic species present in the
Black Sea region, of which five (Caloplaca
arcisproxima,C. austrocitrina,C. communis,
C. confusa, and C. nigromarina) are proposed
here as species new to science. Three speci-
mens were classified as Caloplaca sp. because
they do not fit any recognized species, but
they comprise insufficient material to merit
the description of new species. It seems prob-
able that more species of Caloplaca remain to
be discovered in the Black Sea region.
Above species level, the molecular analysis
revealed four large, internally diverse groups
in the ‘Caloplaca citrina clade’. The first and
most basal comprises C. citrina,C. havaasii,
C. ora p.p., C. maritima and C. communis
(PP = 0·99 and BS = 97). The second
group, formed by C. calcitrapa,C. nigroma-
rina,C. marina,C. microthallina and C. con-
fusa has a rather low support (PP = 0·91,
BS = 37). The third group includes C. flav-
ocitrina and C. geleverjae (PP = 1·00,
BS = 89), while the fourth includes C. di-
chroa,C. austrocitrina,C. limonia,C. arcis and
C. arcisproxima (PP = 0·93, BS = 30). The
backbone of the tree is, however, not strongly
supported, which is not surprising when
using only the ITS sequences.
Morphology
Thallus morphology of the Caloplaca cit-
rina clade is very variable. It contains species
with soredia (47% of the species examined),
blastidia (12%), a granular thallus surface
(12%), and a thallus without vegetative dia-
spores (29%). The size of soredia, blastidia
or granules is a useful key character in some
species (Fig. 4). The thallus is always auto-
trophic and lichenicolous species are prob-
ably absent from this group. It varies from
yellow to orange (only exceptionally grey, in
Caloplaca geleverjae) and may be continuous
or formed of discrete granules, areoles or
squamules, up to 550 µm thick and up to
2·7 mm diam. The true cortex is usually in-
conspicuous, formed of 1–2 layers of cells.
The alveolate cortex (for details see Materials
and Methods) is more distinctly developed,
up to c. 50 µm thick, paraplectenchymatous
and formed of thin-walled cells, (3·0–)
5·4 ± 1·1 (–10·0) µm diam. (n= 479). An
epinecral layer is usually present, but thin, up
to c. 20 µm thick. The algal layer fills most of
the thallus interior and is made up of algal
cells (5·0–) 11·4 ± 3·3 (–32·0) µm diam.
(n= 537) as well as ± isodiametric fungal
cells, (3·0–) 5·2 ± 1·2 (–9·0) µm diam.
(n= 537). The medulla is mostly incon-
spicuous, formed of loose hyphal strands. In
sorediate species, the development of the
soralia differs markedly between species and
provides important key characters. Three
different types are described here (Fig. 2);
the ‘confusa-type’, where blastidia are pro-
duced first at the margins of the areoles and ±
labriform soralia develop after the blastidia
have eroded or have been shed; the
‘flavocitrina-type’, where soredia are pro-
duced directly in well-delimited labriform
or rarely laminal soralia; and the ‘limonia-
type’, where blastidia first build up on the
thallus surface, and true soralia are produced
from cracked blastidia or after the blastidia
have eroded. These three types of develop-
ment are hardly recognizable when the thal-
lus is too young or over mature and forming
entire sorediate crusts. Soredia may be
single, c. 20–60 µm diam., or grouped in
consoredia.
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 577

578 THE LICHENOLOGIST Vol. 41

The apothecia are small, up to 1·6 mm
diam., and sessile. Discs (slightly concave–)
flat to slightly (or strongly) convex, ± uni-
formly orange. The apothecial margins
c. 40–210 µm thick, ± persistent and paler
than the discs, zeorine, formed of a ± proso-
plectenchymatous true exciple and a thalline
exciple, which may be indistinct and hidden
below the true exciple in young apothecia,
but raised and distinct in older ones. The
hypothecium (including subhymenium) is
(30–) 102 ± 31 (–230) µm thick (n= 103),
formed of dense, intricate, translucent hy-
phae. The hymenium is c. 60–90 µm thick
and the asci are of Teloschistes-type, cylin-
drical, (34–) 51 ± 6 (–72) × (7–) 11·5 ± 2
(–18) µm (n= 99). The ascospores are
c. 9–17 × 4–8 µm large and belong to one of
two types: the common, thin-walled ‘citrina-
type’ or the rare, thick-walled ‘sand-clock’
type (Fig. 3). The ascospore septa are
c. 3–7 µm wide. The paraphyses are simple,
branched or anastomosed, c. 1–2·5 µm thick,
with widened tips, up to 7 µm.
The pycnidia are inconspicuous, hardly
recognizable as dots with a different tinge
of yellow or orange, c. 80–130 µm diam.
The conidiophores are variable in height,
branched and anastomosed in parts (Fig. 3).
The conidiogenous cells are variable in shape,
but mostly obtuse triangular, c. 4–6 µm
diam. The conidia are acro- or pleurogenous,
ellipsoid to bacilliform, c. 2–4 × 1–1·5 µm.
Chemistry
All the species treated here contain an-
thraquinones of the chemosyndrome A
(sensu Søchting 1997) with parietin as a
dominant compound, and small amounts of
emodin, fallacinal, parietinic acid and telo-
schistin. This chemosyndrome is the most
common within Teloschistaceae. No signifi-
cant differences were observed among the
species treated (Table 2); some minor differ-
ences are of a quantitative nature only, such
as greater amounts of fallacinal in Caloplaca
arcis,C. austrocitrina,C. flavocitrina and C.
F. 1.Bayesian consensus phylogeny of the Caloplaca citrina group inferred from 105 nuclear ITS sequences. Node
support values (in black) are Bayesian posterior probabilities and numbers in grey represent bootstrap values
obtained after 1000-times resampled parsimony heuristics (bootstrap values under 50% are not shown). Groupings
recovered by parsimony heuristics yet not by the Bayesian inference are depicted by lines to the right of the tree.
F. 2.Development of soralia in the Caloplaca citrina group. A, confusa-type; B, flavocitrina-type; C, limonia-type.
Note that initial and late stages are almost identical in all types. Scales: A–C = 100 µm.
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 579

nigromarina, than in the rest of species
treated. The thallus cortex and the apothecia
are K+ violet, C−, P−, UV± orange. In sec-
tions, the hymenium and the subhymenium
are I+ blue, but the true exciple is I−. Excep-
tionally, in Caloplaca geleverjae, anthraqui-
nones are absent from the K− thallus.
Phenotypic variability and the
taxonomic value of the characters
In most of the characters studied, pro-
nounced phenotypic variability was ob-
served. Some extreme examples of variability
in width of the areoles, size of vegetative
diaspores, width of the exciple, the hy-
menium thickness, width of paraphysis tips
and ascospore characters are shown for
particular species in Fig. 4. In these charac-
ters, there are no, or only small, overlaps in
dimensions between different specimens of
the same species making it difficult to find
diagnostic morphological characters.
A comparison of the species treated using
vegetative diaspore, thallus thickness and
ascospore characters is shown in Fig. 5.
Some species may be grouped, such as Calo-
placa communis,C. geleverjae and C. limonia
F. 3.Caloplaca citrina group. A, C. calcitrapa ascospores, citrina type spores (top row) (CBFS JV5486), sand glass
type spores (bottom row) (CBFS JV6112); B, C. communis, vertical section of a pycnidium (CBFS JV3369). Scales:
A=10µm;B=20µm.
580 THE LICHENOLOGIST Vol. 41

F. 4.Intraspecific variability in selected characters in some Caloplaca species. Distributions of data are expressed as means ± SD (boxes) and extremes. Extremes
depicted when nR10. Sample accession numbers and numbers of measurements are given at the bottom and top, respectively.
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 581

F. 5.Variability in some characters among the Caloplaca species treated. Distributions of data are expressed as
mean ± SD (boxes) and extremes.
582 THE LICHENOLOGIST Vol. 41

T 3.List of species with the Caloplaca citrina morphology occurring on seashore cliffs in the Northern Hemisphere with
data on their distributions and references. Species marked by ** are treated here in detail; those marked by * are treated briefly
under notes on other species
Species † Distribution / ecology References
C. arcis (Poelt & Veˇzda) Arup ** Cc Europe, Mediterranean, the Black Sea
region
Arup 2006a;thispaper
C. arcisproxima Vondrák, Arup &
Søchting sp. nov. ** Cc Black Sea coast, eastern Mediterranean This paper
C. austrocitrina Vondrák, Arup &
Søchting sp. nov. ** Cc Central, eastern and southern Europe;
mostly on concrete
This paper
C. bolacina (Tuck.) Herre Western coast of North America Arup 1993b, 1995b
C. britannica R. Sant. *British Isles; restricted to maritime
rocks
Laundon 1992b
C. calcitrapa Nav.-Ros., Gaya &
Cl. Roux) **
Cc Black Sea coast, Mediterranean; on
calcareous sea-shore rocks, if inland?
Navarro-Rosinés et al.
2000; this paper
C. citrina (Hoffm.) Th. Fr. *Cc Northern and central Europe,
elsewhere not confirmed
Arup 2006a
C. communis Vondrák, Arup &
Søchting sp. nov. ** Cc Black Sea coast, eastern Mediterranean This paper
C. confusa Vondrák, Arup &
Søchting sp. nov. ** Cc Black Sea coast, Mediterranean, Azores This paper
C. coronata (Kremp. ex Körb.) J.
Steiner
Europe; mostly on inland calcareous
rocks
e.g. Foucard 2001
C. dichroa Arup ** Cc Europe; mostly on inland calcareous
rocks
Arup 2006a;thispaper
C. flavocitrina (Nyl.) H. Olivier ** Cc Europe, North America (confirmed
from Hawaii), Asia?; epiphytic or
epilithic, mostly inland species
Arup 2006a; Vondrák et
al. 2007; this paper
C. flavogranulosa Arup Western coast of North America Arup 1993a, 1995b
C. geleverjae Khodosovtsev & S.
Kondr. ** Cc Crimean Peninsula; coastal cliffs Khodosovtsev et al. 2003;
this paper
C. havaasii H. Magn. Cc Norway; known only from type locality,
inland; occurrence on maritime cliffs
not confirmed
Arup 2006c
C. inconnexa (Nyl.) Zahlbr. Mediterranean, Europe; often
lichenicolous, e.g. on Acarospora
cervina
Vondrák et al. 2007
C. inconnexa var. nesodes Poelt &
Nimis (?= C. necator Poelt &
Clauzade)
Black Sea coast, Mediterranean;
lichenicolous on Aspicilia
Nimis & Poelt 1987
C. inconspecta Arup ? Western coast of North America Arup 1995a,b
C. limonia Nimis & Poelt ** Cc Mediterranean, Black Sea region, East
and Central Europe
Nimis et al. 1994; this
paper
C. littorea Tav. ? Atlantic coast of Europe Laundon 1992a; Tavares
1956
C. luteominia (Tuck.) Zahlbr. Western coast of North America Arup 1993b, 1995b
C. ludificans Arup ? Western coast of North America Arup 1995a,b
C. marina (Wedd.) Zahlbr. (syn.:
C. marina subsp. americana
Arup) *
Cc Western coast of North America and
Atlantic coast of Europe
Arup 1992a, 1995b,
1997; Nordin 1972
C. maritima (B. de Lesd.) B. de
Lesd. *Cc Atlantic coast of Europe, West
Mediterranean
Arup 1997
C. microthallina (Wedd.) Zahlbr. *Cc Scandinavian coast, Atlantic coast of
Europe and North America
Arup 1994, 1997b;
Laundon 1992a;
Nordin 1972
C. nigromarina Vondrák, Arup &
Søchting sp. nov. ** Cc Black Sea coast This paper
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 583

that have larger vegetative diaspores, or C.
austrocitrina,C. communis and C. limonia that
have larger areoles, whereas C. dichroa has
somewhat wider ascospores and C. arcis and
C. dichroa have thinner spore septa. How-
ever, the characters are much less useful in a
single sample, which may possess only a part
of a range of variability. For example, speci-
mens of C. limonia from shaded places have
significantly smaller soredia than samples
from sun-exposed rock faces, which produce
blastidia rather than soredia.
Useful key characters are mainly vegeta-
tive, such as thallus colour, presence of dif-
ferent vegetative diaspores of different sizes,
development of soredia and thickness of the
thallus and size of areoles. Apothecial char-
acters (e.g. ascospore size and width of spore
septa) where there is often strong phenotypic
variability and small differences between
species are less useful. Moreover, the soredi-
ate species are rather sparingly fertile and C.
arcisproxima is not known fertile.
Short diagnostic description of Caloplaca
citrina group.
Thallus yellow to orange (except for Calo-
placa geleverjae), continuous or made up of
dispersed granules, areoles or squamules.
Vegetative diaspores, such as soredia and
blastidia are often present. Apothecia small
with orange discs and zeorine margins. True
exciple of elongated cells. Ascospores thin-
walled (except for some specimens in
C. calcitrapa and C. dichroa), with thick
septa.
Unfortunately, these morphological char-
acters are shared with other, phylogenetically
unrelated species, for example, members of
the Caloplaca dolomiticola group. The pos-
ition of each species must therefore be con-
firmed by molecular data. Some superficially
similar species of the Caloplaca lactea group
(e.g. C. crenulatella and C. interfulgens) and of
the C. squamosa group (e.g. C. subsoluta)do
not fit the ‘Caloplaca citrina morphology’.
The former has large, narrow-ellipsoid asco-
spores with thin septa (Navarro-Rosinés &
Hladun 1996) and the latter has a paraplect-
enchymatous true exciple (Arup 1992b;
Wetmore 2003).
Species with the ‘Caloplaca citrina morpho-
logy’ occurring on seashore cliffs are listed
in Table 3. It is noted there that not all of
the species belong to the Caloplaca citrina
clade.
T 3.Continued
Species † Distribution / ecology References
C. ora Poelt & Nimis* Cc Mediterranean Nimis & Poelt 1987
C. phlogina (Ach.) Flag.*Europe; mostly on bark or shrub twigs
but also on concrete close to sea
shore
Arup 2006a; Vondrák et
al. 2007
C. rosei Hasse Western coast of North America Arup 1992a, 1995b
C. ruderum (Malbr.) J. R. Laun-
don
? Great Britain, Continental Europe?; on
mortar and soft limestone in walls, if
on coastal cliffs?
Laundon 1976
C. sorediella Arup Great Britain, coastal rocks Arup 2006b
C. soropelta (Hansen, Poelt &
Søchting) Søchting
Greenland and Svalbard Hansen et al. 1987;
Søchting 1992
C. thamnoblasta Nimis & Poelt Mediterranean; on calcareous sea-shore
rocks
Veˇzda 1993
† column: Cc indicates species belonging to the Caloplaca citrina clade; ? position of species uncertain.
584 THE LICHENOLOGIST Vol. 41

Key to the saxicolous Caloplaca species occurring on coastal cliffs in the Black Sea
region
1 Thallus pale grey, rarely slightly yellow, formed of areoles ± entirely covered by
blastidia. ..............................C. geleverjae
Thallus yellow to orange, in shaded sites pale-yellow to yellow-grey .......2
2(1) Soredia absent, but blastidia may be present ....................3
Soredia present, thallus sometimes blastidiate only, but in well-developed popula-
tions, soredia always present where blastidia have eroded (limonia-type of
soralia) ......................................5
3(2) Vegetative diaspores absent, but large thallus granules may be present ......4
Blastidia present on surface and at margins of inner areoles, (30–) 75 ± 21
(–130) µm, thallus usually with short-lobed margins ..........C. arcis
4(3) Thallus variable, consisting of convex areoles, (90–) 190 ± 75 (–450) µm thick, with
granular surface, or thallus formed by dispersed granules; granules (70–) 180 ± 66
(–380) µm, apothecia usually abundant, large, (0·2–) 0·6 ± 0·2 (–1·3) mm diam.,
often with crenulate margin; common on hard siliceous coastal rocks ......
................................... C. communis
Thallus of flat to convex areoles, only (80–) 129 ± 36 (–210) µm thick; surface
smooth, not granular, on coastal limestone or lime-rich schistose rocks .....
....................................C. calcitrapa
5(2) Thallus thin, up to 200 µm thick, areolate, never squamulose, composed of a greyish-
yellow to yellow sorediate crust only, soredia small (20–) 42 ± 14 (–63) µm;
epiphytic or on loess and concrete; not found on rock .......C. phlogina
Thallus mainly thicker, up to 550 µm thick, soredia on average larger, 23–320 µm, of
confusa,flavocitrina or limonia-type (Fig. 2); known from coastal cliffs .....6
6(5) Soralia ± delimited, developing mainly from margins of areoles (confusa or
flavocitrina-type of soralia), but old thalli may be entirely sorediate ......8
Soralia not delimited, of limonia-type; thallus often entirely sorediate/blastidiate
...........................................7
7(6) Thallus (100–) 248 ± 111 (–550) µm thick, dull to bright yellow, sometimes with
minute marginal lobes, sorediate; soredia coarse, (26–) 85 ± 54 (–320) µm, old
thalli may form a thick, entirely sorediate crust ...........C. limonia
Thallus (70–) 118 ± 36 (–210) µm thick, yellow or orange, without marginal lobes,
areoles ± entirely sorediate/blastidiate, vegetative diaspores smaller, (23–) 40 ± 11
(–61) µm diam., usually both, orange and yellow thallus morphs are present in the
localities ................................C. dichroa
8(6) Thallus of ± umbilicate squamules, with margins divided into minute lobes ....
..................................C. arcisproxima
Thallus areolate or squamulose, but not with margins divided into minute lobes
...........................................9
9(8) Thallus areolate (rarely squamulose), (80–) 192 ± 70 (–380) µm thick, areoles
(squamules) large, (0·25–) 0·74 ± 0·32 (–1·7) mm wide, old thalli usually entirely
covered by soredia; mainly on concrete ............C. austrocitrina
Areoles/squamules smaller, up to 180 µm thick and up to 1·4 mm wide; thallus
usually not forming entire sorediate crust ...................10
(a difficult species, lacking sufficient diagnostic morphological characters)
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 585

10(9) Thallus mostly yellow, soralia of flavocitrina type predominate; if on coast, then
usually on calcareous substrata (concrete, calcareous sandstones, etc.) .....
...................................C. flavocitrina
Thallus mostly yellow-orange (in shaded sites yellow); on siliceous coastal cliffs
..........................................11
11(10) Confusa,flavocitrina and limonia types of soralia present ........C. confusa
Only confusa and flavocitrina types of soralia present .......C. nigromarina
Taxonomy
Caloplaca arcis (Poelt & Veˇzda) Arup
Lichenologist 38: 8 (2006).—Caloplaca citrina var. arcis
Poelt&Veˇzda in Veˇzda, Lichenes Selecti Exsiccati, fasc.
99 (1990); type: Austria, Styria, distr. Feldbach,
Riegersburg, alt. 400 m, on andesite rock, 1990, G.
Kantvilas, H. Mayrhofer & A. Veˇzda (A. Veˇzda: Lich.
Sel. Exsicc. 2470, BM!, PRM!—isotypes).
(Fig. 6A & B)
Thallus yellow, areolate or formed of
tightly aggregated squamules; marginal
areoles have a ± lobate character. Areoles/
squamules (80–) 150 ± 39 (–220) µm thick
(n= 21) and (0·3–) 1·11 ± 0·7 (–3·0) mm
wide (n= 47). Thallus surface smooth or
covered by granules and blastidia; in old
thalli, vegetative diaspores may cover most of
the thallus surface. Granules/blastidia (30–)
75 ± 21 (–130) µm diam. (n= 31). Cortex
indistinct, alveolate cortex (7–) 18 ± 8 (–32)
µm high (n= 18).
Apothecia present in 37% of investigated
specimens, (0·41–) 0·6 ± 0·13 (–0·88) mm
diam. (n= 19). Discs slightly concave to
flat. Exciple (70–) 105 ± 21 (–140) µm thick
(n= 15), zeorine, sometimes crenulate. The
true exciple/thalline exciple ratio very
variable; thalline exciple enlarged with
age. Hymenium 70–100 µm thick (n= 12).
Paraphyses tips widened to (3·0–) 4·4 ± 0·6
(–5·5) µm (n= 27). Ascospores (9·0–) 12·0 ±
1·25 (–14·0) × (4·5–) 5·5 ± 0·9 (–7·5) µm
(n= 20); length/breadth ratio c. 2·2; asco-
spore septa (3·0–) 4·0 ± 0·5 (–5·0) µm thick
(n= 20), c. 0·33 of ascospore length.
Conidia (1·5–) 2·4 ± 0·5 (–3·0) × (1·0–)
1·25 ± 0·25 (–1·5) µm (n= 26).
Remarks. This species is characterized
mainly by its blastidiate thallus with short-
lobed margins. In some cases, it is hardly
distinguishable from Caloplaca limonia, but
in well-developed populations, C. arcis has
distinct marginal lobes, and blastidia (sore-
dia are absent) are restricted to central parts
of the thallus; whereas C. limonia often has an
entirely sorediate/blastidiate thallus surface.
Phylogeny. According to the DNA analy-
ses, Caloplaca arcis forms a well-supported
monophyletic group (PP 1·00, BS = 98)
with two sequences of C. arcisproxima in a
sister position and C. limonia as the second
closest taxon.
Ecology and distribution. Caloplaca arcis
occurs mainly on inland sun-exposed, hard
siliceous, but usually base-rich rock faces.
Although it is also known from pure lime-
stone (Arup 2006a), it was not found on
calcareous substrata in the Black Sea region.
When it occurs on coastal rocks, it usu-
ally avoids the supralittoral zone (Kiten,
Bulgaria, extremely sheltered shore: from
1 m upwards; Sinop, Turkey, extremely ex-
posed shore: from 130 m upwards). Accom-
panying species: Caloplaca aff. furax,C.
teicholyta,C. thracopontica,Candelariella
plumbea,Catillaria chalybeia,Diplotomma
alboatrum s. lat., Lecanora albescens,L. cam-
pestris,Phaeophyscia orbicularis,Verrucaria
macrostoma f. furfuracea, and Xanthoria cf.
calcicola. The species is widely distributed
in Europe. Here, it is newly recorded from
Bulgaria, Italy, the Netherlands, Slovakia,
Turkey and the Canary Islands.
Specimens examined.Bulgaria: Black Sea coast:
Burgas, Tsarevo, Rezovo, 2005, J. Vondrák (CBFS
JV3036); Burgas, Kiten, rocks near mouth of Karaagach
river S of town, 2007, J. Vondrák (CBFS JV6093).—
Italy: Sardinia: Arburese, on volcanic rock alt. 320–
380 m, 1986, W. Brunnbauer (GZU, sub C. citrina).—
586 THE LICHENOLOGIST Vol. 41

The Netherlands: Liemers, 2003, A. Aptroot 59582
(ABL); Zwartemeerdijk, 2004, A. Aptroot 59871
(ABL).—Slovakia: Muránská planina Mt:. in valley
Hrdzavá, a1t. 1220–1270 m, 1999, A. Guttová, J. Halda
& Z. Palice (SAV, sub C. citrina).—Spain: Canary
Islands: Tenerife, Aguamansa, 1986, W. Brunnbauer (W,
sub Caloplaca sp.).—Turkey: Black Sea coast: Sinop, alt.
c. 100 m, 2007, J. Vondrák (CBFS JV5426, 6106).
F. 6.Morphology of Caloplaca species. A & B, Caloplaca arcis;C&D,C. arcisproxima;E&F,C. austrocitrina.
Scales: A–F=1mm.
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 587

Caloplaca arcisproxima Vondrák,
R
ˇíha, Arup & Søchting sp. nov.
Thallus squamulis umbilicatis (0·44–) 0·78 ± 0·21
(–1·18) mm latis contextus, margo squamularum in
lobulos minutos divisus. Soralia typo C. confusae vel
flavocitrinae.
Typus: Ukraine, Crimean Peninsula, Alushta, coastal
rocks SW of Ribachye, 44°45'35.36$N, 34°35'10.30$E,
on supralittoral diabasic rock, 27 May 2007, J. Vondrák
(CBFS JV5473—holotypus; CBFS JV6038, hb. C &
LD—isotypi). ITS sequence of the holotypus:
EU563425.
(Fig. 6C & D)
Thallus yellow to yellow-orange, of solitary
or rarely aggregated, ± umbilicate squam-
ules. Squamules (80–) 129 ± 36 (–240) µm
thick (n= 29) and (0·44–) 0·78 ± 0·21
(–1·18) mm wide (n= 22). Squamules flat,
smooth, with margins divided into minute
lobes. Marginal soralia of confusa or flavo-
citrina type. Soredia/blastidia (25–) 48 ± 16
(–86) µm diam. (n= 26). Cortex indistinct,
alveolate cortex (8–) 24 ± 9 (–46) µm thick
(n= 36).
Apothecia absent in specimens investi-
gated.
Conidia (2·5–) 3·0 ± 0·3 (–4·0) × (0·5–)
1·0 ± 0·25 (–1·5) µm (n= 15).
Etymology.Arcisproxima means close to
arcis. The new species is phylogenetically
close to Caloplaca arcis and the species also
share some morphological characters (e.g.,
distinct squamules).
Remarks. The species is characterized
mainly by its umbilicate squamules, with
margins divided into minute lobes and by the
presence of soralia of confusa or flavocitrina
types. The species may be confused with
Caloplaca confusa,C. flavocitrina or C. nigro-
marina, but it differs in its ± umbilicate
squamules, with somewhat raised margins. It
somewhat resembles C. arcis in its squam-
ules, which are often divided into minute
lobes, but C. arcisproxima differs in the pres-
ence of confusa or flavocitrina types of soralia.
Phylogeny. The monophyly of the two
sequences obtained from Caloplaca arcis-
proxima was not confirmed by the parsimony
analysis (BS = 70%) or the Bayesian infer-
ence. However, this may be due to poor
lineage sorting in a young species or to
the fact that only two specimens have been
analysed. A close affinity to C. arcis is shown
(PP = 1·00), but the two species clearly differ
from each other in their morphology.
Ecology and distribution. Based on the
known localities, the species prefers base-
rich, hard siliceous rocks (e.g. diabasic), but
was found also on ±soft, lime-rich claystone.
It is known only from maritime sites, usually
from the supralittoral zone. Accompanying
species: Caloplaca cf. aegea,C. biatorina s.
lat., Caloplaca aff. furax,C. teicholyta,Candel-
ariella plumbea,C. vitellina,Catillaria chaly-
beia,Diplotomma alboatrum s.lat., Lecanora
albescens,L. dispersa s. lat., Phaeophyscia or-
bicularis,Verrucaria macrostoma f. furfuracea,
and Xanthoria cf. calcicola. The species is so
far known from the Crimean Peninsula in the
Black Sea region and Crete in the eastern
Mediterranean.
Specimens examined: Ukraine: Crimean Peninsula:.
Cape Meganom, 2002, A. Khodosovtsev (KHER 3030,
sub C. citrina).—Greece: Crete: Agios Pavlos, alt.
c. 215–240 m, 1997, H. Mayrhofer (GZU); Ano Vian-
nos, Sidonia, alt 10–50 m, 2005, J. Vondrák (CBFS
JV4125); Mires, Kali Limenes, siliceous rock, alt.
c. 100 m, 2005, J. Vondrák (CBFS JV3877).
Caloplaca austrocitrina Vondrák, R
ˇíha,
Arup & Søchting sp. nov.
Caloplacae citrinae similis, differt thallo areolato (raro
squamuloso) (80–) 192 ± 70 (–380) µm crasso, areo-
lisque (squamulis) majoribus (0·25–) 0·74 ± 0·32 (–1·7)
mm latis. Soralia typo C. flavocitrinae. Praecipue in operi
concreto habitat.
Typus: Ukraine,Crimean Peninsula,. Alushta
(), slopes above sea c. 1 km SW of Semidvorje
(), alt. c. 100 m, 44°42'48.10$N,
034°27'12.10$E, on vertical side of concrete wall, 13
June 2006, J. Vondrák (CBFS JV5236—holotypus;
CBFS JV5285, LD—isotypi). ITS sequence of the
holotypus: EU563419
(Fig. 6E & F)
Thallus yellow, rarely orange (e.g. the
specimen from Crete), areolate or formed
of tightly aggregated squamules. Areoles/
588 THE LICHENOLOGIST Vol. 41

squamules (80–) 192 ± 70 (–380) µm thick
(n= 22) and (0·25–) 0·74 ± 0·32 (–1·7) mm
wide (n= 51). Squamules flat, smooth, with
marginal soralia of flavocitrina type; in old
thalli, soralia may cover the whole thallus
surface. Soredia (22–) 36 ± 10 (–61) µm
diam. (n= 34), sometimes gathered to con-
soredia. Cortex or alveolate cortex developed,
(6–) 19 ± 8 (–40) µm thick (n= 30).
Apothecia present in 50% of the specimens
investigated, (0·32–) 0·46 ± 0·1 (–0·65) mm
diam. (n= 16). Disc in mature apothecia flat
to convex. Exciple (50–) 86 ± 20 (–130) µm
thick (n= 16), zeorine; in young apothecia,
the thalline exciple is hidden below the true
margin while in old apothecia, the thalline
exciple is well-developed and persisting. Hy-
menium 60–80 µm thick (n= 5). Paraphyses
tips widened to (3·0–) 4·4 ± 0·9 (–6·0) µm
(n= 15). Ascospores (8·5–) 11·25 ± 1·5
(–140) × (4·5–) 6·0 ± 0·5 (–6·5) µm (n= 16);
length/breadth ratio c. 1·88. Ascospore septa
(3·0–) 4·5 ± 1·0 (–5·5) µm thick (n= 16),
c. 0·4 of ascospore length.
Conidia (2·0–) 2·8 ± 0·5 (–4·0) × (1·0–)
1·2 ± 0·2 (–1·5) µm (n= 15).
Etymology. The name reflects the distri-
bution of the species in Europe.
Remarks. This species is distinguished by
its thick, areolate thallus (rarely squamu-
lose), its large areoles (squamules), and its
soralia of the flavocitrina type. Old thalli are
usually entirely covered by soredia. Caloplaca
flavocitrina, occurring on the same kind of
substratum (predominantly concrete), dif-
fers in its thinner and smaller squamules.
Phylogeny. Although morphologically
distinguishable, the specimens of C. austro-
citrina do not receive any support as a mono-
phyletic taxon, but this does not preclude
possible monophyly in other genes. More
work on other genes may resolve the phy-
logeny of C. austrocitrina, but the lineage
sorting among other genes may also be poor.
In the phylogenetic analysis C. austrocitrina
groups with C. limonia,C. arcisproxima and
C. arcis (PP = 0·92).
Ecology and distribution. Most of the
records are from lime-rich artificial substrata,
for example, concrete and mortar, but the
species is also known from limestone. It
occurs inland as well as in coastal areas and
does not avoid substrata close to sea level, for
example faces of concrete walls in harbours.
Accompanying species: Caloplaca aurantia,
C. biatorina s. lat., C. crenulatella,Candelari-
ella aurella,Lecania leprosa,Lecanora albes-
cens,L. dispersa,L. muralis,Rinodina pityrea,
Verrucaria macrostoma f. furfuracea, and V.
muralis. The species is probably common in
south-eastern Europe and is also known from
central Europe (Austria, Czech Republic and
Germany) and the European part of Russia.
Selected specimens examined. Austria: Styria: Joglland,
Miesenbach, alt. c. 1040 m, 2003, J. Hafellner (GZU,
sub C. citrina).—Bulgaria: Black Sea coast: Burgas,
Pomorie, 2005, J. Vondrák (CBFS JV4631).—Czech
Republic: South Bohemia:C
ˇeské Budeˇjovice, 2003,
J. Vondrák (CBFS JV991).—Germany: Bavaria:
Munich, 1978, E. Albertshofer & H. Hertel (W, Lich.
Alpium 328, sub C. citrina).—Greece: Crete: Rethimno,
ruin of fort Fortezza, 2005, J. Vondrák (CBFS
JV4195).—Romania: Dobrogea: Romania, Tulcea, near
shore of Razim Lake (Lacul Razim), 2007, J. Vondrák
(CBFS JV6168).—Russia: Black Sea coast: Novoros-
siysk, coastal rocks near Dyurso, 2007, J. Vondrák
(CBFS JV5474, dupl. in C); Sochi, coast 2 km SE of
Loo, 2007, J. Vondrák (CBFS JV6097, 6166).—
Slovakia: Bukovské vrchy hills: Nová Sedlica, 1989, I.
Pišút (SAV, sub C. citrina). Podunajská nížina lowland:
Vrakúnˇ, 1990, I. Pišút (SAV, sub C. citrina).—Ukraine:
Black Sea coast: Odessa, Czernomorskoye, 2007, J.
Vondrák (CBFS JV5476, dupl. in LD).
Caloplaca calcitrapa Nav.-Ros., Gaya
& Cl. Roux
Bull. Soc. linn. Provence 51: 147 (2000); type: France,
Eastern Pyrenees, Languedoc Roussellon, Tautavel, alt.
400–450 m, on calcareous rock (BCC 13385, holoty-
pus). ITS sequence of an isotype: DQ173227.
(Fig. 7A & B)
Thallus yellow, rarely orange, areolate or
bullate, occasionally thin with a rather endo-
lithic character. Snail-grazed forms with flat
areoles are common. Areoles (80–) 129 ±
36 (–210) µm thick (n= 26) and (0·32–)
0·52 ± 0·16 (–0·97) mm wide (n= 20). Veg-
etative diaspores absent. Cortex indistinct,
alveolate (8–) 25 ± 9 (–35) µm thick
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 589

(n= 15); epinecral layer usually with inter-
mixed crystals.
Apothecia frequent, (0·28–) 0·46 ± 0·15
(–0·75) mm diam. (n= 20). Disc flat to
strongly convex in old apothecia. Exciple
(40–) 100 ± 34 (–210) µm thick (n= 57),
zeorine, the true exciple/thalline exciple ratio
very variable; thalline exciple enlarged with
F. 7.Morphology of Caloplaca species. A & B, Caloplaca calcitrapa;C&D,C. communis;E,C. confusa;F,C. dichroa.
Scales: A–F=1mm.
590 THE LICHENOLOGIST Vol. 41

age. Hymenium (60–) 68 ± 6 (–80) µm thick
(n= 15). Paraphyses tips widened to (4·0–)
5·5 ± 0·75 (–7·0) µm (n= 25). Ascospores
(7·0–) 11·0 ± 2·0 (–14·0) × (4·5–) 6·0 ±
0·75 (–7·5) µm (n= 32); length/breadth ratio
c. 1·83. Ascospores usually thin-walled (in the
Black Sea region); thick-walled, sand-clock
spores observed only in two samples. Asco-
spore septa (3·0–) 4·5 ± 1·0 (–6·5) µm thick
(n= 32), c. 0·4 of ascospore length.
Conidia (2·5–) 3·3 ± 0·4 (–4·0) × (1·0–)
1·3 ± 0·25 (–1·5) µm (n= 15).
Remarks. According to Navarro-Rosinés
et al. (2000), the species is characterized
mainly by thick-walled, sand-clock asco-
spores, but these were only rarely observed in
specimens from the Black Sea and Marmara
Sea regions (CBFS JV6109 & JV6112).
More common forms with thin-walled spores
were confirmed by the DNA analysis (CBFS
JV3408, JV5486 & JV6100).
The areolate or bullate thallus of Caloplaca
calcitrapa may be confused with the inland
and facultatively lichenicolous species C.
inconnexa, but C. calcitrapa seems to be ±
maritime and not lichenicolous.
Phylogeny. The four very divergent ITS
sequences of Caloplaca calcitrapa form a
monophyletic group with good PP support
(1·00) but a very poor bootstrap support
(BS = 53) in a sister position to a clade in-
cluding C. confusa,C. marina,C. micro-
thallina, and C. nigromarina (PP = 0·91,
BS < 50).
Ecology and distribution. In the Black Sea
region, the species inhabits strongly calcare-
ous substrata, for example hard limestone
and lime-rich schist. In coastal areas, it
specifically occurs in the supralittoral zone
(Rusalka, Bulgaria, sheltered shore: from
1 m upwards; Kamen Brjag, Bulgaria, ex-
posed shore: from 6 m upwards). In western
Mediterranean, it is also known from inland
localities (Navarro-Rosinés et al. 2000).
Accompanying species: Caloplaca cf. aegea,
C. biatorina s. lat., C. decipiens,C. erythro-
carpa,C. ferrarii,C. limonia,C. navasiana,
Candelariella aurella,Diplotomma alboatrum
s. lat., Lecanora albescens,L. campestris,
L. dispersa,Rinodina gennarii,Verrucaria
nigrescens, and Xanthoria cf. calcicola. The
species is known from Algeria, France, Italy,
Morrocco and Spain (Navarro-Rosinés et al.
2000, Roux et al. 2003). It is recorded here
for the first time from Bulgaria, Turkey and
Ukraine.
Specimens examined.Bulgaria: Black Sea coast:
Kavarna, limestone cliffs on seashore 1·5 km NE of
Kamen Brjag, 2007, J. Vondrák (CBFS JV5486, 6117,
6130).—Turkey: Black Sea coast: Istanbul, Kemerbur-
gaz, Karaburun, 2005, J. Vondrák (CBFS JV3408);
Kandıra, coastal limestone rocks 6 km E of Cebeci,
2007, J. Vondrák (CBFS JV6112); Marmara Sea coast:
Armutlu, coastal rocks 5·5 km SW of Esenköy, 2007,
J. Vondrák (CBFS JV6100, 6109); Bandırma, coastal
rocks near Yenice, 2007, J. Vondrák (CBFS JV6118,
6123, 6125, 6129, 6180); Gallipoli Peninsula, coastal
limestone cliffs 1 km NE of Abide monument, 2007,
J. Vondrák (CBFS JV6103).—Ukraine: Crimean Penin-
sula: Sudak, coastal rocks at W part of Cape Meganom,
2007, J. Vondrák (CBFS JV5906); Sea of Azov coast,
1995, A. Redchenko (KHER 2986, sub C. marina).
Caloplaca communis Vondrák, R
ˇíha,
Arup & Søchting sp. nov.
Caloplaca calcitrapae et maritimae similis, differt super-
ficie thalli grosse granulato granulis (70–) 180 ± 66
(–380) µm latis. Habitat in rupibus maritimis tantum
praecipue silcieis.
Typus: Bulgaria, The Black Sea coast, Burgas,
Tsarevo, coastal rocks SE of town, near small boat-
factory, 42°08'49.7$N, 27°52'48.2$E, on siliceous rock
in supralittoral zone, 30 November 2005, J. Vondrák
(CBFS JV4620—holotypus; isotypi will be distributed
in J. Vondrák: Sel. Exs.Caloplaca, fasc. 2).
(Fig. 7C & D)
Thallus yellow (with orange tint in popula-
tions from Crete) areolate, bullate to
minutely squamulose. Areoles dispersed or
contiguous and then overgrowing each other,
forming thick crusts in luxuriant popula-
tions, (90–) 190 ± 75 (–450) µm thick
(n= 53) and (0·32–) 0·86 ± 0·43 (–2·8) mm
wide (n= 121). When the thallus is well-
developed, its surface is coarsely granular;
granules (70–) 180 ± 66 (–380) µm diam.
(n= 135). Cortex present, (5–) 17 ± 7·5
(–46) µm thick (n= 134).
Apothecia frequent, (0·2–) 0·6 ± 0·2 (–1·3)
mm diam. (n= 97). Disc in mature apoth-
ecia slightly concave to flat, never distinctly
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 591

convex. Exciple (40–) 100 ± 34 (–210) µm
thick (n= 57), often crenulate; true exciple
fills c. ⅓–½ of the exciple width, thalline
exciple prominent. Hymenium (50–) 68 ± 12
(–100) µm thick (n= 48). Paraphyses tips
widened to (3·0–) 5·0 ± 1·0 (–7·0) µm
(n= 80). Ascospores (10·0–) 13·0 ± 1·5
(–18·0) × (4·0–) 6·5 ± 1·0 (–9·0) µm
(n= 78); length/breadth ratio c. 2·0. Asco-
spore septa (3·0–) 5·0 ± 1·0 (–7·0) µm thick
(n= 78), c. 0·4 of ascospore length.
Conidia (2·0–) 2·8 ± 0·3 (–3·5) × (1·0–)
1·2 ± 0·2 (–2·0) µm (n= 52).
Etymology. The name reflects a common
occurrence of this species on siliceous sea-
shore cliffs.
Remarks. The yellow thallus, covered by
coarse granules, together with its occurrence
restricted to coastal habitats (mainly sili-
ceous rocks), are the main characters dis-
tinguishing Caloplaca communis from the
similar C. calcitrapa and C. inconnexa. Never-
theless, some morphotypes of C. communis
have a thallus with an indistinctly granular
surface and if such a morphotype occurs on a
calcareous substratum then it is hardly dis-
tinguishable from the likewise coastal C.
calcitrapa. When the areoles/squamules are
dispersed, C. communis may resemble the
North and West European C. microthallina,
but this species has smaller squamules, 0·3–
0·5(–1·0) mm wide (Arup 1994). Morpho-
types with a more compact thallus and a less
distinct granular surface resemble C. mar-
itima, which is not confirmed from the re-
gion. Caloplaca ora, which is a heterogenous
taxon, is another similar ‘species’, but with
a typically orange-red thallus without a
granular surface.
Phylogeny. All the sequences sampled of
C. communis except one form a monophyletic
group in the molecular analysis but with low
support (PP = 0·87, BS < 50)). The position
of the remaining sequence assigned to this
species (EU563409) is controversial, appear-
ing in a polytomy between C. communis,
C. maritima and C. ora p. p. As in the case of
C. austrocitrina, sampling of more loci could
possibly make parts of the molecular tree
more consistent. The genetic affinity of
C. maritima and C. ora p. p. to C. communis
is, however rather strong (PP = 0·99,
BS = 78).
Ecology and distribution.Caloplaca commu-
nis is one of the commonest species in the
lichen communities of the lower supralittoral
zone on hard siliceous rocks. Only very
rarely, it occurs on sandstone, claystone and
limestone cliffs. The species is restricted to
a narrow zone close to sea level; sheltered
shores: 2–4 m (Tuapse, Russia), 1–2 m
(Kiten, Bulgaria); exposed shores: 3–7 m
(Sarp, Turkey), 4–17 m (Bulgaria, Rezovo)
and 4–19 m (Sinop, Turkey). In the eastern
Mediterranean, where the salinity is higher
and the climate warmer and drier, the species
may occur higher above sea level: c. 20–50 m
(Agia Pelagia, northern Crete) and c. 100 m
(Kali Limenes, southern Crete). Accompa-
nying species: Amandinea punctata,Caloplaca
cf. aegea,C. confusa,C. fuscoatroides,C. limo-
nia,Lecanora campestris,Rinodina confragosa,
R. gennarii. It is widely distributed in the
Black Sea region, Marmara Sea and in the
eastern Mediterranean. It is known from
Bulgaria, Greece, Italy, Russia, Turkey and
Ukraine.
Specimens examined:Bulgaria: Black Sea coast:
Burgas, Kiten, 2007, J. Vondrák (CBFS JV6088);
Burgas, Sozopol, 2007, J. Vondrák (CBFS JV6121);
Burgas, Tsarevo, 2005, J. Vondrák (CBFS JV3369);
Burgas, Tsarevo, Rezovo, 2005, J. Vondrák (CBFS
JV3043).—Greece: Crete: Ano Viannos, Sidonia, 2005,
J. Vondrák (CBFS JV4114); Iraklio, Agia Pelagia, 2005,
J. Vondrák (CBFS JV3803, 3836); Mires, Kali Limenes,
2005, J. Vondrák (CBFS JV3763).—Italy: Sardinia:
Planargia, 1983, W. Brunnbauer (W, sub C. micro-
thallina).—Russia: Black Sea coast: Tuapse, coastal
rocks NW of town, Tuapse, 2007, J. Vondrák (CBFS
JV6108). Sea of Azov coast: Taman Peninsula, clay coast
4·5 km NW of Kuchugury, 2007, J. Vondrák (CBFS
JV6104).—Turkey: Black Sea coast: Amasra, coastal
rocks near Çakrazboz, 2007, J. Vondrák (CBFS
JV6127); Cide, coastal rocks near Denizkonak, 2007,
J. Vondrák (CBFS JV6115); Istanbul, Kemerburgaz,
Kilyos, 2005, J. Vondrák (CBFS JV3367, 3467, 3471,
3472); Kandıra, sand dunes and coastal limestone rocks
6 km E of Cebeci, 2007, J. Vondrák (CBFS JV6119);
Lülenburgaz, Demirköy, Limanköy, 2005, J. Vondrák
(CBFS JV3037); Lülenburgaz, Vize, Kiyiköy, 2005,
J. Vondrák (CBFS JV3042); Ordu, coastal rocks near
Mersin, 2007, J. Vondrák (CBFS JV6128); Zonguldak,
592 THE LICHENOLOGIST Vol. 41

coastal rocks near Ilıksu, 2007, J. Vondrák (CBFS
JV6116). Marmara Sea coast: Armutlu, coastal rocks
5·5 km SW of Esenköy, 2007, J. Vondrák (CBFS
JV6114); Karabiga, 2007, J. Vondrák (CBFS JV6110,
6111, 6124).—Ukraine: Crimean Peninsula: Cape
Alushtinska miska, 2000, A. Khodosovtsev (KHER 2979,
sub C. microthallina); Cape Meganom, 1999 & 2002, A.
Khodosovtsev (KHER 2977, 2980, 2981, sub C. micro-
thallina); Cape Plaka, 1999, A. Khodosovtsev (KHER
2982, sub C. microthallina); Karadag, 2000, A. Khodos-
ovtsev (KHER 2976, sub C. microthallina).
Caloplaca confusa Vondrák, R
ˇíha,
Arup & Søchting sp. nov.
Caloplaca flavocitrinae et nigromarinae similis, differt
praesentia soraliorum typo C. confusae,flavocitrinae et
limoniae. Habitat in rupibus maritimis silcieis tantum.
Typus: Bulgaria, Black Sea coast, Burgas, Kiten,
rocks near mouth of Karaagach river S of town,
42°13'31.26$N, 27°46'40.46$E, on coastal, base-rich,
siliceous rock, 9 April 2007, J. Vondrák (CBFS
JV6206—holotypus; BM, C, GZU, LD—isotypi). ITS
sequence of the holotypus: EU563449.
(Fig. 7E)
Thallus yellow to yellow-orange, of dis-
persed squamules, or continuous, areolate.
Areoles/squamules (80–) 121 ± 28 (–180) µm
thick (n= 16) and (0·2–) 065 ± 0·3 (–1·4)
mm
wide (n= 34). Areoles and squamules
flat, smooth or covered by blastidia, with
laminal or marginal soralia of confusa,flavoc-
itrina or limonia type; soredia (18–) 31 ± 9
(–43) µm diam. (n= 28), sometimes gath-
ered to small consoredia. Cortex poorly
developed; alveolate cortex (7–) 18 ± 8·5
(–40) µm thick (n= 15).
Apothecia infrequent (25% of samples
fertile), c. 04–1·0 mm in size (n= 7). Discs
observed in mature apothecia ± flat. Exciple
50–110 µm thick (n= 7), zeorine, but ap-
pearing biatorine; in young apothecia, the
true exciple fills the whole exciple width, as
the thalline exciple is hidden below the true
exciple. Hymenium 70–80 µm thick (n= 9).
Paraphyses tips widened to (3·5–) 4·5 ±
0·5 (–5·5) µm (n= 18). Ascospores (10·0–)
11·5 ± 1·0 (–13·0) × (4·5–) 5·5 ± 1·0 (–7·0)
µm (n= 17); length/breadth ratio c. 2·1. As-
cospore septa (3·5–) 5·0 ± 1·0 (–7·0) µm
thick (n= 17), c. 0·43 of ascospore length.
Conidia c. 3–3·5 × 1–1·5 µm.
Etymology. The name expresses the con-
fusing position of the new species in the
phylogenetic tree; although very similar to
Caloplaca flavocitrina and C. nigromarina,itis
not closely related to them.
Remarks. This species may be character-
ized by its squamulose to areolate thallus
with all three types of soralia (confusa,flavo-
citrina and limonia types), but in some cases,
it is indistinguishable from Caloplaca flavo-
citrina and C. nigromarina.
Phylogeny. The sequences of Caloplaca
confusa form a monophyletic group, but the
support is not very strong (PP = 0·89,
BS = 46). However, it is morphologically
distinct from its genetically closest relative,
C. microthallina, in the analyses. The support
for this sister relationship is not very strong
(PP = 0·88, BS = 47).
Ecology and distribution. Caloplaca confusa
is restricted to hard siliceous, mainly vol-
canic, sea shore cliffs in the supralittoral
zone; from c. 2 m upwards in sheltered
shores and 5–18 m on an exposed shore
(Sinop, Turkey). Accompanying species:
Caloplaca cf. aegea,C. communis,C. cf. holo-
carpa,C. limonia,C. thracopontica,Catillaria
chalybeia,Hyperphyscia adglutinata,Lecania
cf. aipospila,Lecanora campestris,L. dispersa,
L. cf. salina,Rinodina gennarii, and Xanthoria
cf. calcicola.
It is probably widely distributed in the
Black Sea region and the Mediterranean, and
is also known from the Atlantic coast.
Records from Azores, Bulgaria, France and
Italy are confirmed by molecular data. There
are also collections from Georgia, Russia,
Turkey and Ukraine, but these were not con-
firmed by molecular data and have been
omitted from this study.
Specimens confirmed by ITS data. Bulgaria: Black Sea
coast: Burgas, Tsarevo, coastal rocks between Tsarevo
and Ahtopol, 2005, J. Vondrák (CBFS JV3435).—
France: Corsica: Ajaccio, on volcanic rock in alt. 50 m,
1993, J. Hafellner (GZU, sub C. citrina).—Italy: Sicily:
Isole Pelagie, Linosa, 1992, J. Poelt (GZU, sub C.
citrina).—Portugal: Azores: Sao Jorje, 1992, F. Berger
(hb. Berger).
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 593

Caloplaca dichroa Arup
Lichenologist 38: 13 (2006); type: Sweden, Västergöt-
land, Klefva infra Mösseberg. In saxis calcaris, 1914,
Vrang. Malme: Lichenes suecici exsiccati 525 (LD—
holotype).
(Fig. 7F)
Thallus areolate, of two colour forms, yel-
low and orange. Areoles (70–) 118 ± 36
(–210) µm thick (n= 19) and (0·24–) 0·6 ±
0.·3 (–1.·2) mm wide (n= 20), ± entirely
covered by blastidia/soredia of limonia type,
(23–) 40 ± 11 (–61) µm diam. (n= 30). Cor-
tex indistinct, composed of 1–2 layers of cells;
a thicker alveolate cortex developed in spots.
Apothecia frequent, (0·42–) 0·49 ± 0·06
(–0·64) mm (n= 15). Disc in mature apoth-
ecia slightly concave to slightly convex.
Exciple (50–) 83 ± 18 (–110) µm thick (n=
15), zeorine, the true exciple/thalline exciple
ratio very variable; thalline exciple often
covered by blastidia/soredia, enlarged with
age. Hymenium c. 70–80 µm thick (n= 5).
Paraphyses tips swollen up to (2·5–) 4·5 ± 1·0
(–6·0) µm (n= 34). Ascospores thick-walled
(walls up to 2 µm) or thin-walled (walls less
than 0·5 µm), (11·0–) 14·0 ± 1·5 (–16·0) ×
(6·0–) 7·25 ± 0·75 (–9·0) µm (n= 15);
length / width ratio c. 1·93. Ascospore septa
(3·5–) 4·0 ± 0·4 (–4·5) µm thick (n = 15),
c. 0·29 of ascospore length.
Conidia (1·5–) 2·5 ± 0·5 (–3·0) × (1·0–)
1·25 ± 0·25 (–1·5) µm (n= 20).
Remarks. The species is characterized by
its sorediate/blastidiate thallus surface, limo-
nia type of soralia and thin thallus without
marginal lobes. The yellow and orange thal-
lus forms, often growing side by side, are also
diagnostic. It can be confused with Caloplaca
austrocitrina and C. limonia, but the former
has a generally thicker thallus and a different
type of soralium, and the latter has a thicker
thallus, larger vegetative diaspores and lacks
the orange morphotype. For differences from
C. citrina and C. coronata see Arup (2006a).
The thick-walled ascospores seem to be diag-
nostic in the Nordic coutries (Arup 2006a),
but in the Black Sea region some specimens
have thin-walled spores. A similar pattern
has also been observed in some British speci-
mens.
Phylogeny.Caloplaca dichroa forms one of
the best-supported clades in our analysis; it
has full support (PP = 1·00, BS = 100). This
branch appears as a sister taxon to a clade of
C. arcis, C. arcisproxima and C. limonia with
fairly good support in the Bayesian analysis
(PP = 0·93).
Ecology and distribution. Although this
species is known also from concrete and mor-
tar (Arup 2006a; Vondrák et al. 2007), in the
Black Sea region, it has only been collected
from ± sun-exposed, hard or soft calcareous
stones or cliffs. It is a typical inland species
avoiding maritime conditions; on coastal
cliffs it grows from 6 m upwards on sheltered
shores (Rusalka, Bulgaria) and from 16 m
and upwards on more exposed shores
(Kamen Brjag, Bulgaria). Accompanying
species: Candelariella aurella,C. rhodax,
Lecanora albescens,L. dispersa,Phaeophyscia
nigricans,P. orbicularis,Physcia adscendens,
Physconia grisea,Verrucaria macrostoma f.
furfuracea, and V. nigrescens s. lat.
Selected specimens examined.Austria: Styria: Eisen-
erzer Alpen, Kammern im Liesingtal, 1997, J.&A.
Hafellner (GZU, sub C. citrina); Grazer Bergland,
Mixnitz, alt. 1620 m, 2005, J. Hafellner (GZU, sub C.
citrina); Fischbacher Alpen, Rettenegg, 2002, J. Hafell-
ner (GZU, sub C. citrina). Tirolia: Hohe Tauern Mts,
Matrei, alt. 1000 m, 1998, J. Hafellner (GZU, sub
C. citrina).—Bulgaria: Kavarna, Cape Kaliakra, 2007,
J. Vondrák (CBFS JV6177).—Great Britain: England:
V. C. 34, West Gloucestershire: Bristol, rocks on right
side of Avon River near Clifton suspension bridge, 2006,
J. Vondrák (CBFS JV4155).—Hungary: Mt Bakony:
Hárskut, 1968, K. Verseghy (BP 75193, 75204).—
Romania: Dobrogea: Jurilovca, rocky cliff at Dolos¸man
Cape, 2007, J. Vondrák (CBFS JV5324, 5337); Tulcea,
Popina Island (Insula Popina) in Razim Lake (Lacul
Razim), 2007, J. Vondrák (CBFS JV6178, 6179).—
Russia: Sea of Azov coast: Taman Peninsula, on soft
limestone, 4·5 km NW of Kuchugury, 2007, J. Vondrák
(CBFS JV6176).—Ukraine: Crimean Peninsula: Feodo-
sia, Karadag, near village Kurortnoye, 2007, J. Vondrák
(CBFS JV5477).
Caloplaca flavocitrina (Nyl.) H. Olivier
Lich. d’Europe 2: 110 (1908–1910).
Lecanora flavocitrina Nyl., Flora 69: 461 (1886); type:
Great Britain, Staveley, 1886, Martindale (H-Nyl.!, lec-
totype selected by Wade 1965).
594 THE LICHENOLOGIST Vol. 41

(Fig. 8A)
Thallus yellow, of dispersed squamules,
rarely continuous, areolate. Squamules
(70–) 96 ± 17 (–140) µm thick (n= 16) and
(0·12–) 0·4 ± 0·17 (–0·75) mm wide
(n= 60). Squamules flat, smooth, with
marginal soralia of flavocitrina type, but the
soralia sometimes expand and cover the
complete surface of the squamules. Soredia
(16–) 29 ± 9 (–65) µm diam. (n= 56), some-
times gathered into consoredia. Cortex thin,
F. 8.Morphology of Caloplaca species. A, Caloplaca flavocitrina;B,C. geleverjae;C&D,C. limonia;E,C.
nigromarina;F,C. phlogina. Scales: A–F=1mm.
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 595

usually of 1–2 layers of cells; alveolate cortex
(4–) 9 ± 5 (–25) µm thick (n= 44). A white
or yellow fibrillar prothallus was observed on
smooth substrata.
Apothecia infrequent (30% of samples fer-
tile), (0·3–) 0·43 ± 0·1 (–0·75) mm diam.
(n= 24). Disc in mature apothecia slightly
concave to flat. Exciple (70–) 85 ± 13
(–110) µm thick (n= 16), zeorine, the true
exciple/thalline exciple ratio very variable;
thalline exciple enlarged with age. Hymenium
(65–) 75 ± 6 (–80) µm thick (n= 14). Para-
physes tips swollen to (3·0–) 5·0 ± 1·0 (–7·0)
µm (n= 32). Ascospores (10·0–) 11·5 ±
1·25 (–14·5) × (4·0–) 5·5 ± 1·0 (–7·0) µm
(n= 27); length/width ratio c. 2·1. Ascospore
septa (3·0–) 4·5 ± 1·0 (–7·0) µm thick
(n= 27), c. 0·39 of ascospore length.
Conidia (2·0–) 3·25 ± 0·5 (–4·0) × (1·0–)
1·15 ± 0·25 (–1·5) µm (n= 14).
Remarks. This species has a squamulose
to areolate thallus with mainly flavocitrina
type of soralium. It is hardly distinguishable
from C. confusa and C. nigromarina, but its
thallus is usually yellow and the flavocitrina
type of soralium persists. It differs also in its
ecology, being mainly an inland species, and
is rather rare on sea-shore rocks.
Phylogeny. The sequences of Caloplaca
flavocitrina form a well-supported mono-
phyletic group in the Bayesian analysis
(PP = 0·99), but the parsimonious analysis
gave a weaker support (BS = 61). Together
with C. geleverjae and an undescribed taxon,
represented by the sequence EU563389, it
forms a well supported clade (PP = 1·00,
BS = 89), that appears as a sister group to a
large clade with, for example, C. dichroa and
C. arcis, but with rather poor support
(PP = 0·84, BS < 50).
Ecology and distribution.Caloplaca flavo-
citrina is one of the ecologically broadest
species in the C. citrina group; it occurs on
bark, wood, concrete and on a variety of
rocks. In maritime conditions, it is mainly
restricted to concrete or calcareous rocks.
It is confirmed from siliceous coastal cliffs
only in the most humid part of the Black
Sea region in Georgia and NE Turkey.
Accompanying species in coastal localities:
Caloplaca albolutescens,Candelariella aurella,
Diplotomma alboatrum s. lat., Lecanora dis-
persa,Rinodina gennarii, and Verrucaria
macrostoma f. furfuracea.
As it is known throughout Europe and
from the Hawaiian Islands, it may have
a circumpolar distribution. It is a new record
for Georgia, Italy, Russia, and Turkey.
Specimens confirmed by ITS nrDNA data.Bulgaria:
Black Sea coast: Burgas, Pomorie, on concrete, 2005, J.
Vondrák (CBFS JV3425); Burgas, Tsarevo, Sinemorec,
epiphytic, 2004, J. Vondrák (CBFS JV2106).—
Georgia: Adjara: Batumi, valley of river Acharistskali
near Shuakhevi, 2007, J. Vondrák (CBFS JV6094);
Batumi, rocks near coast on S periphery of Gonio, 2007,
J. Vondrák (CBFS JV6098).—Italy: Livorno, volcanic
coastal rock, alt. 20–50 m, 1993, J. Hafellner (GZU,
sub C. citrina).—Russia: Black Sea coast: Gelendzhik,
coastal rocks W of Krinitsa (near Betta), 2007, J.
Vondrák (CBFS JV6089); Sochi, coast 2 km SE of Loo,
2007, J. Vondrák (CBFS JV6095).—Turkey: Black
Sea coast: Rize, coastal outcrops close to city, 2007,
J. Vondrák (CBFS JV6090).—USA: Hawaii: Maui,
Haleakala, volcanic rock, alt. 1700 m, 1996, C. Wetmore
(GZU, sub C. citrina).
Caloplaca geleverjae Khodosovtsev & S.
Kondr.
Ukr. Botan. Journ.60: 294 (2003); type: Ukraine,
Crimea AR, Feodosia region, cape Meganom, on con-
glomerates in supralittoral zone, 2002, A. Khodosovtsev
(KW—holotype; KHER!, CBFS!—isotypes). ITS
sequence of a topotype: EU563423.
(Fig. 8B)
Thallus pale grey, rarely yellowish (mainly
at apothecial primordia), consisting of dis-
persed or continuous areoles, areolate.
Areoles (110–) 147 ± 18 (–180) µm thick
(n= 15) and (0·34–) 0·63 ± 0·16 (–0·87)
mm wide (n= 15). Squamules ± entirely
covered by blastidia, (48–) 85 ± 31 (–148)
µm diam. (n= 15); true soredia absent. Cor-
tex indistinct; alveolate cortex (18–) 28 ± 9
(–50) µm thick (n= 15).
Apothecia frequent in the known popu-
lation, (0·37–) 0·48 ± 0·07 (–0·62) mm
diam. (n= 15). Disc in mature apothecia
slightly concave to flat. Exciple (70–) 93 ± 14
(–110) µm thick (n= 14), zeorine, but thal-
line exciple hidden below the true exciple in
596 THE LICHENOLOGIST Vol. 41

young apothecia. Hymenium c. 60–80 µm
thick (n= 5). Paraphyses tips swollen to
(5·0–) 6·0 ± 0·75 (–7·0) µm (n= 15). As-
cospores (10·0–) 13·25 ± 1·5 (–15·0) ×
(5·0–) 6·5 ± 1·0 (–8·0) µm (n= 15); length/
breadth ratio c. 2·04. Ascospore septa (3·0–)
4·75 ± 0·75 (–6·0) µm thick (n= 15),
c. 0·36 of ascospore length.
Conidia (2·5–) 3·0 ± 0·4 (–3·5) × (1·0–)
1·3 ± 0·2 (–1·5) µm (n= 15).
Remarks. The predominantly greyish thal-
lus colour distinguishes Caloplaca geleverjae
from other related species. The blastidiate
species, Caloplaca thamnoblasta, is somewhat
similar in thallus structure, but its thallus
is usually yellow and it does not belong to
the C. citrina group (J. Vondrák & J. Šoun,
unpublished molecular data).
Phylogeny.Caloplaca geleverjae (one se-
quence included in the analysis) is a sister
species to C. flavocitrina (PP = 0·98;
BS = 60).
Ecology and distribution. Caloplaca gele-
verjae is known only from the type locality in
the Crimean Peninsula, where it occurs on ±
shaded, base-rich conglomerate in the lower
supralittoral zone, at altitudes below 10 m.
The accompanying species in the type local-
ity were few and included for example, Calo-
placa limonia and C. cf. nigromarina.
Specimen examined.Ukraine: Crimean Peninsula:
Cape Meganom, 2007, J. Vondrák & A. Khodosovtsev
(CBFS JV5415—topotype).
Caloplaca limonia Nimis & Poelt
in Nimis, Poelt, Tretiach, Ottonello, Puntillo & Veˇzda,
Bull. Soc. Linn. Provence 45: 252 (1994); type: Italy,
Isole Egadi, Marettimo, 1991, J. Poelt (GZU!—
holotype). ITS sequence of a paratype: EU563467.
(Fig. 8C & D)
Thallus dull to bright yellow, often white-
pruinose, forming a continuous areolate
crust. Areoles/squamules (100–) 248 ± 111
(–550) µm thick (n= 44) and (0·18–) 0·98 ±
0·51 (–2·6) mm wide (n= 140). Areoles and
squamules flat, covered by blastidia or by
laminal soralia of limonia type; soredia/
blastidia (26–) 85 ± 54 (–320) µm diam.
(n= 138). Cortex/alveolate cortex (4–) 15 ±
10 (–55) µm thick (n= 109).
Apothecia frequent, c. (0·25–) 0·6 ± 0·25
(–1·6) mm diam. (n= 114). Disc in mature
apothecia slightly concave to flat. Exciple
(70–) 121 ± 31 (–200) µm thick (n= 25),
zeorine, thalline exciple prevailing in old
apothecia, sometimes covered by blastidia/
isidia. Hymenium (60–) 78 ± 17 (–130) µm
thick (n= 50). Paraphyses tips widened to
(3·0–) 4·5 ± 0·5 (–6·0) µm (n= 72). Asco-
spores (9·0–) 13·0 ± 1·5 (–16·0) × (3·0–)
6·5 ± 1·0 (–9·0) µm (n= 77); length/breadth
ratio c. 2·0. Ascospore septa (3·0–) 5·5 ± 1·0
(–8·0) µm thick (n= 77), c. 0·42 of ascospore
length.
Conidia (2·5–) 3·5 ± 0·8 (–5·5) × (1·0–)
1·15 ± 0·25 (–1·5) µm (n= 33).
Remarks. The diagnostic characters of this
species are large soredia/blastidia, limonia
type of soralium, ± pale yellow thallus and a
thick apothecial margin often covered by
blastidia/isidia. Caloplaca citrina, that has a
more northern distribution, is probably the
most similar species, but it differs on average
in the smaller vegetative diaspores, which are
often blastidia and not soredia.
Phylogeny. The sequences of Caloplaca
limonia form a well-supported monophyletic
group (PP = 1·00, BS = 94). It appears to be
related to C. arcis along with C. arcisproxima
(PP = 0·80, BS < 50) and more distantly to
C. austrocitrina (PP = 0·92, BS < 50).
Ecology and distribution. Caloplaca limonia
occurs mainly on coastal calcareous rocks or
on base-rich, hard siliceous cliffs in dry sun-
exposed to shaded and damp situations, but
also on twigs of maritime shrubs or on soil. It
is also known from inland localities, but it is
often abundant in maritime habitats, where it
occurs on sheltered shores from 2 m upwards
(Rusalka, Bulgaria); on exposed shores from
20 m (Sinop, Turkey) and 21 m (Rezovo,
Bulgaria). Accompanying species: Caloplaca
cf. aegea,C. albolutescens,C. communis,C.
confusa,C. flavescens,Caloplaca thracopontica,
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 597

Catillaria chalybeia,Diploicia canescens,
Diplotomma alboatrum s. lat., Lecanora albes-
cens,L. dispersa,Physcia adscendens,Rinodina
gennarii,R. pityrea,Verrucaria macrostoma f.
furfuracea, and V. nigrescens s. lat.
Caloplaca limonia is so far reported from
Greece (Sipman & Raus 2002), Italy (e.g.
Nimis et al. 1994) and Ukraine (Khodosovtsev
2001), but it is a common species in the
Mediterranean and the Black Sea region.
New record for Bulgaria, Croatia, Czech
Republic, Georgia, Morrocco, Romania,
Russia, and Turkey.
Selected specimens examined.Bulgaria: Black Sea
coast: Kavarna, limestone cliffs on seashore 1·5 km NE
of Kamen Brjag, 2007, J. Vondrák (CBFS JV6138).
—Croatia: Island of Arbe: Arbe, 1927, J. B. Kümmerte
(BP 27347).—Czech Republic: South Bohemia:Pı´sek,
in town, 2005, J. Vondrák (CBFS JV2515).—Georgia:
Black Sea coast: Batumi, Sarpi (Georgian-Turkish bor-
der), 2007, J. Vondrák (CBFS JV6150).—Italy: Distr.
Verona, Massalongo (BP 27353, ex hb. A. Massal.).—
Morocco: Rabat, 1930, Andreánszky (BP 46517).—
Romania: Dobrogea. Tulcea, Jurilovca, rocky cliff at
Dolos¸man Cape, 2007, J. Vondrák (CBFS JV5376).—
Russia: Black Sea coast: Tuapse, coastal rocks NW of
town, 2007, J. Vondrák (CBFS JV6160).—Turkey:
Black Sea coast: Sinop, coastal rocks on E coast of
peninsula, 2007, J. Vondrák (CBFS JV6143). Aegean
Sea coast: Gallipoli Peninsula, coastal rocks near
Güneyli, 2007, J. Vondrák (CBFS JV6148). Marmara
Sea coast: Karabiga, S-exposed coastal rocks on head-
land E of town, 2007, J. Vondrák (CBFS JV6147).—
Ukraine: Crimean Peninsula: Sudak, coastal rocks at W
part of Cape Meganom, 2007, J. Vondrák (CBFS
JV6019).
Caloplaca nigromarina Vondrák, R
ˇíha,
Arup & Søchting sp. nov.
Similis Caloplaca flavocitrinae, differt squamulis
plerumque laetius flavo-aurantiacis; a C. confusae differt
praesentia soraliorum typo C. confusae et flavocitrinae
tantum. Habitat in rupibus maritimis silcieis tantum.
Typus: Georgia,Adjar,. Sarpi (Georgian-Turkish
border), coastal cliffs N of village, 41°31'58.19$N,
41°32'58.35$E, on siliceous rock, 25 April 2007,
J. Vondrák (CBFS JV6091—holotypus). ITS sequence
of the holotypus: EU563436.
(Fig. 8E)
Thallus yellow-orange (in shaded sites
yellow), of dispersed granules/squamules,
or thallus continuous, areolate. Areoles/
squamules (60–) 106 ± 25 (–170) µm thick
(n= 39) and (0·18–) 0·44 ± 0·19 (–1·25)
mm wide (n= 89). Areoles and squamules
flat, smooth, with marginal soralia of confusa
or flavocitrina type, soredia/consoredia (16–)
31 ± 9 (–50) µm diam. (n= 89). Cortex
present, (5–) 17 ± 8 (–37) µm thick (n= 90).
Apothecia infrequent (40% of specimens
fertile), (0·45–) 0·6 ± 0·1 (–0·8) mm diam.
(n= 15). Disc in mature apothecia flat to
slightly convex. Exciple (40–) 75 ± 33 (–150)
µm thick (n= 15), zeorine; true exciple fills
c. ½ of the exciple width. In young apothecia,
thalline exciple may be hidden below the true
exciple. Hymenium (50–) 70 ± 10 (–90) µm
thick (n= 13). Paraphyses tips widened
to (3·0–) 4·5 ± 1·0 (–6·5) µm (n= 29). Asco-
spores (9·0–) 11·5 ± 1·5 (–15·0) × (4·0–)
6·0 ± 1·0 (–8·5) µm (n= 17); length/
breadth ratio c. 1·9. Ascospore septa (3·5–)
5·0 ± 1·0 (–7·0) µm thick (n= 17), c. 0·43 of
ascospore length.
Conidia (2·0–) 3·2 ± 0·5 (–4·5) × (1·0–)
1·25 ± 0·25 (–1·5) µm (n= 29).
Etymology. The name is derived from the
distribution of the new species on shores of
the Black Sea.
Phylogeny. The sequences of Caloplaca
nigromarina form a fairly well-supported
monophyletic group (PP = 0·94, BS = 66).
At the base of this clade there is a single
specimen (EU563433) that cannot be as-
signed to any known species, but it is appa-
rantly related to C. nigromarina (PP = 1·0,
BS = 84). However, it differs clearly in its
morphology. These two taxa form a sister
group to a clade consisting of C. confusa,C.
marina and C. microthallina (PP = 0·99,
BS = 97).
Remarks.Caloplaca nigromarina shares the
squamulose thallus and marginal soralia with
the very similar species Caloplaca confusa and
C. flavocitrina. However, from the former
species, it differs in the flavocitrina type of
soralia, and from the latter, in the yellow-
orange thallus and in ecology.
Ecology and distribution. This species is
restricted to siliceous seashore cliffs in the
lower supralittoral zone. Accompanying
598 THE LICHENOLOGIST Vol. 41

species are usually few only: for example,
Caloplaca cf. aegea,C. fuscoatroides,Lecanora
albescens, and L. dispersa. The species is so far
known only from the Bulgarian, Georgian
and Turkish Black and Aegean Sea coasts.
Specimens examined.Bulgaria: Black Sea coast:
Burgas, Tsarevo, Sinemorec, coastal rocks, 2004, J.
Vondrák (CBFS JV4983).—Turkey: Aegean Sea coast:
Gallipoli Peninsula, coastal rocks near Güneyli, 2007,
J. Vondrák (CBFS JV5482); Black Sea coast: Istanbul,
Kemerburgaz, Kilyos, 2005, J. Vondrák (CBFS
JV3354); Lülenburgaz, Vize, Kiyiköy, coastal rocks,
2005, J. Vondrák (CBFS JV3035); Zonguldag, Sazköy,
1992, V. John & E. Sauer (GZU, Lich. Anatol. Exsicc.
80, sub C. citrina).
Notes on other species with
Caloplaca citrina morphology
Caloplaca britannica R. Sant.
in Laundon, Lichenologist 24: 2 (1992); type: Great
Britain, Caithness (UPS—holotype; GZU—isotype!).
This taxon was recently misunderstood by
Aptroot & van Herk (2004), who considered
Caloplaca britannica conspecific with C. limo-
nia, that is widely distributed in coastal as
well as inland Europe; also in the Black Sea
region: Crimea (Khodosovtsev 2001, sub
C. limonia). However, C. britannica is a
blastidiate-isidiate species that does not be-
long to the C. citrina clade (U. Arup, un-
published) and is restricted to coastal
cliffs. We investigated two samples from
the Netherlands, named C. britannica by A.
Aptroot, and both specimens belong to
C. arcis.
Caloplaca citrina (Hoffm.) Th. Fr.
Acta Soc. Regiae Sci. Upsal. 3: 218 (1860).—Verrucaria
citrina Hoffm. Deutschlands Flora 198 (1796); type:
Lecanora citrina, Svecia (H, neotype selected by Nordin
1972).
According to the GBIF web site (www.
gbif.org), this species is considered to be
cosmopolitan. However, according to mol-
ecular data (Arup 2006a) and this study,
Caloplaca citrina may be a species with a
rather restricted distribution. The species has
so far been confirmed only from North and
Central Europe. In the Mediterranean and
the Black Sea region it is replaced by other
species such as C. limonia.
Caloplaca marina (Wedd.) Zahlbr.
in Du Rietz, Zur Methodologischen Grundlage der Mod-
ernen Pflanzenzociologie 170 (1921).—Lecanora marina
Wedd., Mém. Soc. Nat. Sc. Cherb.19: 275 (1875); type:
France, Vendée, 1875, Weddell (PC—lectotype selected
by Nordin 1972; TUR, hb.Vainio 7188—isotypes).
This morphologically distinct species has
not been confirmed from the Black Sea
region and is probably not present there.
Reports from Turkey (John & Breuss 2004)
and Ukraine (e.g. Redchenko 2002) are
probably incorrect. The Turkish material
was not revised by us, but the Ukrainian
specimens in KHER (collected by A.
Khodosovtsev or A. Redchenko) were ident-
ified as Caloplaca calcitrapa or C. irrubescens
(Arnold) Zahlbr.
Caloplaca maritima (B. de Lesd.) B. de
Lesd.
Rev. Bryol. Lichénol.22, fasc. 3-4:313 (1953).—
Caloplaca citrina var. maritima B. de Lesd., Sched. ad
Krypt. exsicc., Cent. 17 (1909); type: France, Dun-
kerque, on calcareous stone, B. de Lesdain (Krypt. exsicc.
1667, W!—lectotype selected by Arup 1997a).
Based on molecular data, this species is
closely related to Caloplaca communis.
Whereas, C. communis occurs in the Black
Sea region and the eastern Mediterranean,
C. maritima is probably restricted to the
Atlantic coast of Europe. Samples from
southern France named C. maritima by Arup
(1997a), require confirmation. Some speci-
mens of C. communis with a smooth thallus
without granules are morphologically indis-
tinguishable from well-developed material
of C. maritima from the Atlantic coast of
Europe.
Caloplaca microthallina (Wedd.)
Zahlbr.
Cat. Lich. Univ.7: 247 (1931).—Lecanora microthallina
Wedd., Mém. Soc. Nat. Sc. Cherb. 19: 276 (1875); type:
France, Vendée, 1875, Weddell (PC—lectotype selected
by Nordin 1972).
2009 Caloplaca citrina group in the Black Sea region—Vondrák et al. 599

This species may resemble morphologi-
cally some morphotypes of Caloplaca commu-
nis with minute areoles, but C. microthallina
has not been confirmed from the Black Sea
region by molecular data and it is probably
absent there. Records from Bulgaria (Veˇzda
1975) and Ukraine (e.g. Redchenko 2002)
are based on C. communis and the record
from Turkish Eastern Mediterranean (Pišút
& Guttová 2008) needs revision.
Caloplaca ora Poelt & Nimis
in Nimis & Poelt, Studia Geobotanica 7: 70 (1987); type:
France, Corse, Ajaccio, on granite coastal cliff, 1969,
Lambinon, Rondon & Veˇzda (A. Veˇzda: Lich. Sel. Exs.
849, hb. A. Veˇzda—holotype, W!—isotype).
Based on the samples from GZU and W,
we consider this Mediterranean taxon to
be heterogeneous. Various specimens with
orange, areolate to bullate thalli, otherwise
similar to Caloplaca maritima, are placed
here. However, at least some specimens
named as C. ora, belong to the C. citrina
group (Fig. 1). This taxon requires further
study.
Caloplaca phlogina (Ach.) Flag.
Mém. Soc. Emulat. Doubs: 250 (1886).—Parmelia citrina
var. phlogina Ach., Methodus: 180 (1803); type: Sweden.
Skåne: Lund, on Ulmus, 2005, Arup L05001 (LD!—
neotype selected by Arup 2006a).
(Fig. 8F)
Although very similar to Caloplaca citrina,
this species does not belong to the Caloplaca
citrina clade. The species, known as an epi-
phyte from northern Europe (Arup 2006a)
and northern France (Sérusiaux et al. 1999),
also occurs in the Black Sea region as an
epiphyte, but also on loess soil and on
concrete.
Specimens confirmed by ITS nrDNA data.Romania:
Constanta, Mangalia, on concrete, 2005, Vondrák
(CBFS JV3437, sub C. citrina, GenBank nr
EU563460).—Russia: Taman Peninsula: on loess and
on Salicornia shrubs, 2007, Vondrák (CBFS JV6060,
6061, 6224).
General results from the phylogenetic
analysis
While this is principally a taxonomic study
the results can be considered in a broader
context. Some general conclusions and ob-
servations are discussed below.
Cryptic or semi-cryptic species.
The results show that there were several
unrecognized species among the taxa ana-
lyzed. Some of these, for example, C. commu-
nis and C. arcisproxima, can be fairly easily
characterized morphologically even if they
are similar to other species of the group.
Others, such as C. austrocitrina and C. nigro-
marina, are very difficult to distinguish mor-
phologically from C. flavocitrina and many
forms of these species are indistinguishable
from each other without molecular analyses.
Morphological characters may provide only a
hint as to which species the specimen be-
longs, but there is such a large overlap that
these characters are very hard to use. These
difficult species are more or less cryptic, but
we propose to call them ‘semi-cryptic’, since
there are differences in their ecology and
distribution and we feel that there will be a
significant loss of information if they are not
described.
In addition, the species which are difficult
to separate by morphology are not sister
species. For example, C. austrocitrina,C.
nigromarina and C. flavocitrina belong to
three different phylogenetic clades (Fig. 1),
mixed with several other species, which are
clearly non-cryptic. If one were to lump the
similar species into single taxa this would
produce polyphyletic taxa, and that would
not be acceptable. Therefore, we feel that the
semi-cryptic species also need to be formally
described. Another example of indistinguish-
able taxa is C. citrina and C. phlogina, which
sometimes have identical morphology but
belong to two separate larger clades.
Cryptic species problems have been re-
cently referred to in several works; probably
the best known are studies on phylogenetic
species bounderies in Letharia (Arnerup et al.
600 THE LICHENOLOGIST Vol. 41

2004; Kroken & Taylor 2001). Several ‘cryp-
tic species’ were revealed in Letharia, but
some of the species differ slightly in morpho-
logy, chemistry and distribution, and could
be considered to be ‘semi-cryptic’. Similarly,
the new species in Parmelina quercina s. lat.
(Argüello et al. 2007) or in Parmelia saxatilis
s. lat. (Crespo et al. 2002) could also be
considered to be semi-cryptic. In Caloplaca,
for example, C. albopruinosa and C. alociza
could be considered semi-cryptic, as they are
largely indistinguishable on morphological
grounds (Muggia et al. 2008).
Convergences
There are strong convergences in morpho-
logy in distantly related sorediate species.
For example C. flavogranulosa,C. phlogina,
C. sorediella and C. soropelta are morphologi-
cally very similar to some members of the
C. citrina clade, but belong to different, un-
related groups. It is therefore sometimes not
possible to assign a species to the C. citrina
clade without molecular analyses. Even
within this clade there are very similar species
placed in distant positions of the phylo-
genetic tree (Fig. 1).
Changes in vegetative characters
It is surprising how frequent shifts from a
sorediate to a non-sorediate state or switches
between the different kinds of soralium
(flavocitrina-type vs. limonia-type) seem to
have taken place. Whether these shifts are
due to mutations or variations in the expres-
sion of genes is not known, but it does cause
problems in the understanding of the tax-
onomy. If we assume that the C. citrina clade
evolved from a non-sorediate ancestor, there
has been, for instance a single appearance of
soredia followed by four independent losses
of soredia.
Biodiversity
The number of species in the C. citrina
clade will probably increase if this group is
surveyed in other geographical areas using
molecular data. This is also confirmed by
new data from the British Isles (U. Arup,
unpublished data).
Species distribution
Some taxa, which were considered to be
wide-spread or even cosmopolitan, have in
fact very restricted natural ranges. For in-
stance, C. citrina s. str. is absent from the
Black Sea region and its range may be re-
stricted to only parts of Europe. This same
pattern appears to occur for some maritime
species, for example, Caloplaca marina,C.
maritima, and C. microthallina, which are
absent from the Black Sea region and very
probably from the eastern Mediterranean.
Appraisal of taxonomic importance of
some morphological characters
Our molecular data also suggest that some
ascospore characters may be of less impor-
tance taxonomically than previously thought,
for example, the “sand glass ascospores” with
thick walls, which were used as a diagnostic
character for C. calcitrapa (Navarro-Rosinés
et al. 2000) and C. dichroa (Arup 2006a).
In some specimens of both species, con-
firmed by molecular data, only thin-walled
“citrina-type ascospores” were observed.
This observation is particularly important
taxonomically and further study of the
phenomenon is planned.
We are grateful to André Aptroot for providing some
herbarium material, Pavel Hrouzek (Trˇebonˇ, CZ) for
analyzing the anthraquinones, Per Lassen (Lund) for
providing Latin diagnoses and Jaroslav Šoun (C
ˇeské
Budeˇjovice) for his help with the molecular laboratory
work. Our work was financially supported by the Grant
Agency of the Academy of Sciences of the Czech Repub-
lic (KJB 601410701) and a grant from the European
Commission’s (FP 6) Integrated Infrastructure Initia-
tive programme SYNTHESYS (DK-TAF).
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Accepted for publication 30 March 2009
604 THE LICHENOLOGIST Vol. 41