The Lichenologist 43(2): 113–135 (2011)
doi:10.1017/S0024282910000721
© British Lichen Society, 2011
Taxonomy and phylogeny of the Caloplaca cerina group in
Europe
Jaroslav ŠOUN, Jan VONDRÁK, Ulrik SØCHTING,
Pavel HROUZEK, Alexander KHODOSOVTSEV and Ulf ARUP
Abstract: Using ITS nrDNA sequence data, the Caloplaca cerina group (Teloschistaceae) is defined
here as a monophyletic, but internally richly branched lineage. The group is also characterized by a
combination of morphological and anatomical characters. Its internal lineages are supported by
phenotypic characters in addition to ecology and distribution. Within the large C. cerina group, we
have found at least 20 phylospecies in the temperate zone of the Northern Hemisphere. Two species
complexes do not produce any vegetative diaspores: the polyphyletic, corticolous Caloplaca cerina s. lat.
(six separated cryptic or semi-cryptic species) and the monophyletic C. stillicidiorum s. lat. that grows
mainly on plant debris, small shrubs and bryophytes and consists of at least four internal lineages. All
lineages producing vegetative diaspores (soredia, blastidia, isidia or lobules) are phenotypically
characteristic and represent fairly easily distinguishable species: C. chlorina, C. isidiigera, C. monacensis,
C. subalpina, C. thracopontica, C. turkuensis and C. ulmorum. Only the North American sorediate C.
pinicola possibly represents an aggregate of species. Caloplaca sterilis is described as a new species. A key
to the phenotypically distinguishable species is provided. Lectotypes are designated for C. albolutea,
Caloplaca cerina f. coronulata and for C. monacensis. The Australian C. hanneshertelii belongs to this
group, but this and other possible species from the Southern Hemisphere are not treated here in detail.
Some species traditionally placed in the C. cerina group due to their similar morphology are excluded
here on the basis of our phenotype examinations and molecular data. Caloplaca albolutea, C. mydalaea
and C. virescens are uncertain taxa and their identities still remain unclear.
Key words: cryptic species, key, lichens, lichen-forming fungi, nrDNA ITS, semi-cryptic species,
Teloschistales
Introduction
The Caloplaca cerina group is treated here as
a monophyletic group of species around C.
cerina (Hedw.) Th. Fr. It is morphologically
characterized by a crustose thallus, which is
J. Šoun and J. Vondrák: Department of Botany, Faculty
of Science, University of South Bohemia, Branišovská
31, CZ-370 05, České Budějovice, Czech Republic.
Email: jasoun@centrum.cz
U. Søchting: Section for Ecology and Evolution,
Department of Biology, University of Copenhagen,
Universitetsparken 15, DK-2100 Copenhagen Ø,
Denmark.
P. Hrouzek: Institute of Microbiology, ASCR, Department of Autotrophic Microorganisms, Opatovický
mlýn, CZ-379 81 Třeboň, Czech Republic.
A. Khodosovtsev: Department of Botany, Kherson State
University, 40 Rokiv Zovtnya 27, 7300 Kherson,
Ukraine.
U. Arup: Botanical Museum, Lund University, Östra
Vallgatan 18, SE-223 61 Lund, Sweden.
dark grey to white, devoid of anthraquinones
but containing the grey pigment Sedifoliagrey (K+ violet in section). The apothecia
can be classified as lecanorine with yellow
to orange discs containing non-chlorinated
anthraquinones, even though there is a
strongly reduced true exciple containing
anthraquinones in all species. The thalline
exciple is distinct, persistent and in shades of
grey, devoid of anthraquinones and with a
conspicuous cortex in the lower part. The
ascospores are c. 10–16 m long, c. 6–9 m
wide and have a septum c. 4–7 m thick; the
conidia are bacilliform, 3–4 m long and
c. 1 m wide.
The large morphological variation within
the group (e.g. in type of vegetative
diaspores, disc colour, presence of pruina,
size of apothecia) has resulted in many
specific and infraspecific names described
114
THE LICHENOLOGIST
Vol. 43
T 1. List of species that belong to or have formerly been included in the Caloplaca cerina group by Clauzade & Roux
(1985) and Wetmore (2007). Only the infraspecific names of C. cerina with examined type material are listed. Cce –
affiliation to the C. cerina monophyletic group according our ITS-sequence data; when brackets are used, the placement is
based on phenotype characters only. Names representing accepted species within the C. cerina monophyletic group are in bold
Name
Cce
C. albolutea (Nyl.) Zahlbr.
(Y)
C. amabilis (Fink) Zahlbr.
C. aractina (Fr.) Häyrén
N
N
C. areolata (Zahlbr.) Clauzade
N
C. asserigena (J. Lahm) H. Olivier
N
C. borealis (Vain.) Poelt
N
C. cerina (Hedw.) Th. Fr.
C. cerina s. lat.
Y
Y
C. cerina var. chloroleuca (Sm.) Th. Fr.
C. cerina f. coronulata Th. Fr.
C. cerina var. muscorum (A. Massal.) Jatta
C. cerinoides (Anzi) Yatta
C. chlorina (Flot.) H. Olivier
C. chlorina var. cyanopolia (Nyl.) H. Olivier
C. chlorina var. paradoxa (Vain.) H. Olivier
C. conglomerata (Bagl.) Jatta
C. congrediens J. Steiner
Y
Y
Y
?
Y
Y
Y
N
N
C. conversa (Kremp.) Jatta
N
C. crocea (Kremp.) Hafellner & Poelt
(N)
C. erythrantha (Tuck.) Zahlbr.
(N)
C. gilva (Hoffm.) Zahlbr.
C. gilvolutea (Nyl.) Jatta
C. grimmiae (Nyl.) H. Olivier
Y
?
N
C. hanneshertelii S.Y. Kondr. & Kärnefelt
C. haematites (Chaub. ex St.-Amans) Zwackh
Y
N
C. isidiigera Vězda
C. jemtlandica H. Magn.
C. jemtlandica var. cerinosora E. S. Hansen,
Poelt & Søchting
C. monacensis (Leder.) Lettau
C. muscorum (A. Massal.) M. Choisy & Werner
C. mydalaea (Körb.) Zahlbr.
Y
Y
Y
Y
Y
(Y)
C. pellodella (Nyl.) Hasse
N
C. pinicola H. Magn.
Y
Note
Relationship uncertain, probably belonging to
C. stillicidiorum s. lat.
Synonym to C. pellodella
Differs from the C. cerina group by a distinct true
exciple (often with blackish outer pigmentation)
and conidia c. 4–5 m long
Differs from the C. cerina group by a distinct true
exciple, presence of chlorinated anthraquinones,
etc.
Differs from the C. cerina group by a distinct true
exciple and different anthraquinone content
Differs from the C. cerina group by a distinct true
exciple (often with dark outer pigmentation)
Included in C. cerina s. lat.
Polyphyletic taxon including at least six putative
cryptic or semi-cryptic species
Included in C. stillicidiorum s. lat.
Here synonymized with C. turkuensis
Included in C. stillicidiorum s. lat.
Relationship uncertain
Here synonymized with C. chlorina
Here synonymized with C. chlorina
Perhaps synonym to C. pellodella
Differs from the C. cerina group e.g. by brown
thallus
Differs from the C. cerina group by dark apothecia
and a distinct true exciple (often with blackish
outer pigmentation)
Differs from the C. cerina group e.g. by 3-locular
ascospores
Differs from the C. cerina group e.g. by red pycnidia
containing anthraquinones
Included in C. cerina s. lat.
Relationship uncertain, its identity unclear
Differs from the C. cerina group e.g. by brown
thallus
Australian species, not treated here in detail
Differs from the C. cerina group by a distinct true
exciple (sometimes with dark grey outer
pigmentation) and c. 4–5 m long conidia
Here synonymized with C. turkuensis
Here synonymized with C. turkuensis
Included in C. stillicidiorum s. lat.
Relationship uncertain, probably belonging to
C. stillicidiorum s. lat.
Differs from the C. cerina group by a distinct true
exciple and a distinctly squamulose thallus
2011
The Caloplaca cerina group in Europe—Šoun et al.
Name
Cce
C. pyracea (Ach.) Th. Fr.
N
C. spalatensis Zahlbr.
N
C. squamuloisidiata van den Boom & V. J. Rico
?
C. stillicidiorum (Vahl) Lynge
C. stillicidiorum s. lat.
Y
Y
C. sterilis Šoun, Khodosovtsev & Vondrák
C. subalpina Vondrák, Šoun & Palice
Y
Y
C. thracopontica Vondrák & Šoun
Y
C. ulmorum (Fink) Fink
C. turkuensis (Vain.) Zahlbr.
C. virescens (Sm.) Coppins
C. xerica Poelt & Vězda
Y
Y
(Y)
N
from all over the world (Table 1). Caloplaca
cerina itself is an important species to understand as it is the type for the genus Caloplaca.
The group name has usually been applied
in a broader morphological sense (Clauzade
& Roux 1985; Hansen et al. 1987; Wetmore
1996, 1997, 2004, 2007a), but our unpublished molecular data and phenotype evaluation show that some species commonly
placed in this group belong elsewhere
(Table 1) and they are not treated in detail
here. The first data about the phylogeny of
the group was published by Vondrák et al.
(2008), who described two new species, C.
subalpina Vondrák, Šoun & Palice and C.
thracopontica Vondrák & Šoun. The present
study deals with the phylogeny of the entire
group in Europe and also, to some extent,
North America and western parts of Asia.
Material and Methods
Our study is mainly based on material from the temperate zone of Europe, North America and Asia. Samples
from ASU, B, BM, C, CBFS, GZU, H, KHER, LD,
MIN, PRM, TUR, UPS and the private herbaria of
115
Note
Differs from the C. cerina group by a distinct true
exciple, belonging to the C. holocarpa group
Differs from the C. cerina group by a distinct true
exciple, presence of chlorinated anthraquinones,
etc.
Species with unsettled phylogenetic position;
phenotypically fits the C. cerina group
Included in C. stillicidiorum s. lat.
Monophyletic taxon with at least four internal
phylospecies
A new species described here
Described in a previous paper (Vondrák et al.
2008), not treated here in detail
Described in a previous paper (Vondrák et al.
2008), not treated here in detail
Relationship uncertain, its identity unclear
Differs from the C. cerina group by a distinct true
exciple, presence of chlorinated anthraquinones,
etc.
P. van den Boom, J. Malı́ček, Z. Palice, T. Spribille
and the first author have been used. Morphological
data were taken only from sequenced specimens listed
in Table 2. The specimens were examined with light
microscopy. Anatomical features were measured on
hand-cut sections or squash preparations mounted in
water. Morphological characters were measured on dry
material using a dissecting microscope (×45). Measurements are recorded as (min.–) x ± SD (–max.), where
x = mean value, SD = standard deviation and min./max.
= extremes. The total number of measurements (n) is
given in brackets. Only the ascospores with welldeveloped septa (loculi connected by a thin and distinct
cytoplasmatic channel) were measured.
Molecular markers, DNA extraction, PCR
amplification and sequencing
Initially, we tried to use two loci, mtSSU and nrITS,
for molecular analysis. However, the former proved to
be too conserved for our study, thus we continued only
with the ITS gene, which has suitable variability. Direct
PCR was used for PCR-amplification of the ITS regions
including the 5.8S gene of the nuclear rDNA following
Arup (2006). Primers for amplification were ITS1F
(Gardes & Bruns 1993) and ITS4 (White et al. 1990).
PCR cycling parameters followed Ekman (2001). Products were cleaned using JETquick PCR purification
Spin Kit (Genomed). Both complementary strands
were sequenced either by Macrogen Inc., Korea or by
the Laboratory of Genomics, Biology Centre of the
Academy of Sciences of the Czech Republic.
116
THE LICHENOLOGIST
Vol. 43
T 2. Voucher specimens and GenBank accession numbers of the ITS sequences used in the phylogenetic analysis.
Specimens in bold were already at hand and the remainder were newly produced in this study
Species
C. aurantia
C. flavescens
C. cerina 1 ‘A’
C. cerina 2 ‘A’
C. cerina 3 ‘A’
C. cerina 4 ‘A’
C. cerina 5 ‘A’
C. cerina 6 ‘A’
C. cerina 7 ‘A’
C. sp.*
C. subalpina 1
C. subalpina 2
C. subalpina 3
C. cerina 8 ‘B’
C. cerina 9 ‘B’
C. cerina 10 ‘B’
C. cerina 11 ‘C’
C. cerina 12 ‘C’
C. cerina 13 ‘C’
C. turkuensis 1
C. turkuensis 2
C. turkuensis 3
C. turkuensis 4
C. turkuensis 5
C. turkuensis 6
C. hanneshertelii*
C. isidiigera 1
C. isidiigera 2
C. isidiigera 3
C. isidiigera 4
C. isidiigera 5
C. isidiigera 6
C. isidiigera 7
C. isidiigera 8
C. isidiigera 9
C. isidiigera 10
C. ulmorum 1*
C. ulmorum 2*
C. ulmorum 3
C. monacensis 1
C. monacensis 2
C. monacensis 3*
C. monacensis 4*
C. monacensis 5*
C. monacensis 6*
C. monacensis 7*
C. monacensis 8*
C. monacensis 9
C. monacensis 10*
C. monacensis 11
C. monacensis 12
C. monacensis 13*
Specimen
GenBank accession
number
Spain, Gaya et al. 13681 (BCC)
Spain, Gaya 368 (BCN)
Greece, Vondrák JV4061 (CBFS)
Greece, Vondrák JV3862 (CBFS)
Greece, Vondrák JV4035 (CBFS)
Greece, Vondrák JV3794 (CBFS)
Turkey, Šoun 250 (hb. Šoun)
Spain, Vondrák JV7619 (CBFS)
Greece, Vondrák JV3824 (CBFS)
Tasmania, Frödén 1246 (LD)
Spain, Vondrák JV692 (CBFS)
Czech Republic, Palice 6983 (hb. Palice)
Ukraine, Vondrák JV6072 (CBFS)
Iran, Søchting 11063 (C)
Iran, Šoun 253 (hb. Šoun)
Iran, Søchting 11150 (C)
Iran, Šoun 252 (hb. Šoun)
Sweden, Arup L03347 (LD)
Norway, Arup L02355 (LD)
Sweden, Arup L04229 (LD)
Czech Republic, P. & B. v. d. Boom 33926 (hb. v. d. Boom)
USA, Washington, Spribille 15393 (hb. Spribille)
Czech Republic, Malíček & Palice 1908 (hb. Malı́ček)
Italy, Thor 19355 (UPS)
Czech Republic, Palice et al. s. n. (hb. Palice)
Australia, Kondratyuk 20415 (LD)
Austria, Türk RT32349 (GZU)
USA, Montana, Spribille 12617 (hb. Spribille)
Sweden, Arup L04227 (LD)
Ukraine, Vondrák JV6073 (CBFS)
Slovakia, Šoun 283 (hb. Šoun)
Sweden, Arup L04228 (LD)
Sweden, Arup L02352 (LD)
Switzerland, v. d. Boom 17968 (hb. v. d. Boom)
Austria, Arup L97333 (LD)
Austria, Hafellner & Muggia JH68548 (GZU)
USA, South Dakota, Lich. East. North Am. Exs. 268 (GZU)
USA, Minnesota, Wetmore 90256 (GZU)
USA, Missouri, Wetmore 69226 (LD)
Spain, P. & B. v.d. Boom 38821 (hb. v. d. Boom)
Bulgaria, Vondrák JV3236 (CBFS)
Bulgaria, Vondrák JV762 (CBFS)
Bulgaria, Vondrák JV2244 (CBFS)
Hungary, Vondrák & Khodosovtsev JV6393 (CBFS)
Bulgaria, Vondrák JV2127 (CBFS)
Slovakia, Bayerová et al. 8875 (hb. Palice)
Slovakia, Bayerová et al. 6150 (hb. Palice)
Bulgaria, Vondrák JV2026 (CBFS)
Czech Republic, Vondrák 164 (hb. Šoun)
Ukraine, Khodosovtsev 3042 (KHER)
Slovakia, Bayerová et al. 5366 (hb. Palice)
Czech Republic, Vondrák JV548 (CBFS)
AY233219
EU639601
HM538546
HM538548
HM538547
HM538542
HM538543
HM538544
HM538545
HM538484
EU365854
EU365853
EU365855
HM538539
HM538541
HM538540
HM538485
EU365861
HM538486
HM538490
HM538489
HM538487
HM538491
HM538488
HM538492
HM538483
HM538532
HM538533
EU365856
EU365857
HM538536
HM538538
HM538537
HM538534
AF353959
HM538535
HM538509
HM538510
HM538511
HM538493
HM538494
HM538495
HM538503
HM538505
HM538504
HM538506
HM538508
HM538507
HM538496
HM538498
HM538497
HM538499
2011
The Caloplaca cerina group in Europe—Šoun et al.
117
T 2. Continued
Species
C. monacensis 14*
C. monacensis 15*
C. monacensis 16*
C. aff. thracopontica*
C. sterilis 1*
C. sterilis 2
C. sterilis 3*
C. sterilis 4*
C. thracopontica 1
C. thracopontica 2*
C. thracopontica 3*
C. thracopontica 4*
C. thracopontica 5
C. thracopontica 6*
C. thracopontica 7*
C. chlorina 1
C. chlorina 2
C. chlorina 3
C. chlorina 4
C. chlorina 5
C. chlorina 6
C. chlorina 7
C. chlorina 8
C. chlorina 9
C. chlorina 10
C. chlorina 11
C. chlorina 12
C. chlorina 13
C. pinicola*
C. aff. pinicola 1
C. aff. pinicola 2*
C. aff. pinicola 3*
C. cerina 14 ‘D1’
C. cerina 15 ‘D1’
C. cerina 16 ‘D1’
C. cerina 17 ‘D2’
C. cerina 18 ‘D2’
C. cerina 19 ‘D3’
C. cerina 20 ‘D3’
C. cerina 21 ‘D3’
C. cerina 22 ‘D3’
C. stillicidiorum 1
C. stillicidiorum 2 ‘1’
C. stillicidiorum 3 ‘1’
C. stillicidiorum 4 ‘1’
C. stillicidiorum 5 ‘1’
C. stillicidiorum 6 ‘1’
C. stillicidiorum 7 ‘1’
C. stillicidiorum 8 ‘1’
C. stillicidiorum 9 ‘1’
C. stillicidiorum 10 ‘1’
C. stillicidiorum 11 ‘1’
C. stillicidiorum 12 ‘1’
Specimen
Slovakia, Palice 85 (hb. Palice)
Slovakia, Bayerová et al. 6154 (hb. Palice)
Czech Republic, Vondrák & Šoun JV2885 (CBFS)
Greece, Sipman & Raus 54369 (B)
Romania, Vondrák JV6963 (CBFS) – apothecium
Romania, Vondrák JV6963 (CBFS) – soredia
Romania, Vondrák JV6955 (CBFS)
Bulgaria, Vondrák JV6954 (CBFS)
Bulgaria, Vondrák JV3420 (CBFS)
Bulgaria, Vondrák Sel. Exs. Caloplaca 15 (CBFS)
Bulgaria, Vondrák JV3419 (CBFS)
Turkey, Šoun 301 (hb. Šoun)
Turkey, Šoun 302 (hb. Šoun)
Turkey, Vondrák JV5419 (CBFS)
Turkey, Šoun 363 (hb. Šoun)
Denmark, Søchting 9233 (C)
USA, Montana, Spribille 11895 (hb. Spribille)
Czech Republic, Vondrák JV1292 (CBFS)
Canada, Søchting 7086 (C)
Bulgaria, Vondrák JV2055 (CBFS)
Bulgaria, Vondrák JV2226 (CBFS)
Czech Republic, Vondrák JV2056 (CBFS)
Czech Republic, Vondrák JV1294 (CBFS)
Czech Republic, Vondrák JV3034 (CBFS)
Belgium, P. & B. v. d. Boom 30644 (hb. v.d. Boom)
Czech Republic, Vondrák JV2982 (CBFS)
Czech Republic, Vondrák JV3120 (CBFS)
Austria, Vondrák JV1879 (CBFS)
USA, North Dakota, Wetmore 80302 (GZU)
USA, Montana, Spribille & Arvidson 4731 (hb. Spribille)
Canada, British Columbia, Spribille 18340 (hb. Spribille)
USA, Montana, Spribille 9397 (hb. Spribille)
Finland, Palice 11906 (hb. Palice)
Czech Republic, Vondrák JV2316 (CBFS)
Sweden, Foucard s. n. (LD)
Greece, Vondrák JV3776 (CBFS)
Greece, Spribille 16209 (hb. Spribille)
Slovakia, Palice 9009 (hb. Palice)
Norway, Søchting 10045 (C)
Austria, Hafellner JH70279 (GZU)
Spain, Vondrák JV2548 (CBFS)
Spain, Vondrák JV6287 (CBFS)
Austria, Hafellner JH62279 (GZU)
Greece, Vondrák JV4054 (CBFS)
Austria, J. & A. Hafellner JH52814 (GZU)
France, Gueidan s. n. (BCN)
Austria, Hafellner & Muggia JH67075 (GZU)
Spain, P. & B. v. d. Boom 27071 (hb. v. d. Boom)
Hungary, Šoun 61 (hb. Šoun)
Czech Republic, Halda & Palice 11720 (hb. Palice)
Bulgaria, Vondrák Sel. Exs. Caloplaca 12 (CBFS)
Ukraine, Smerychyns’ka 0401(4)3 (KW)
Hungary, Vondrák & Šoun JV4382 (CBFS)
GenBank accession
number
HM538500
HM538501
HM538502
HM538523
HM538528
HM538531
HM538530
HM538529
HM538524
EU365846
EU365847
HM538525
HM538527
EU365848
HM538526
HM538512
HM538513
HM538521
HM538522
EU365859
HM538514
HM538517
HM538516
HM538515
HM538518
HM538520
EU365858
HM538519
HM538479
HM538480
HM538482
HM538481
HM538476
HM538475
AF353958
HM538477
HM538478
HM538471
HM538472
HM538473
HM538474
HM538470
HM538468
HM538469
HM538466
EU639607
HM538467
HM538461
HM538462
HM538463
EU365860
EU681284
HM538465
118
THE LICHENOLOGIST
Vol. 43
T 2. Continued
Species
C. stillicidiorum 13 ‘1’
C. stillicidiorum 14 ‘2’
C. stillicidiorum 15* ‘2’
C. stillicidiorum 16* ‘2’
C. stillicidiorum 17 ‘2’
C. stillicidiorum 18 ‘2’
C. stillicidiorum 19* ‘2’
C. stillicidiorum 20 ‘3’
C. stillicidiorum 21 ‘3’
C. stillicidiorum 22 ‘3’
C. stillicidiorum 23 ‘3’
C. stillicidiorum 24 ‘4’
C. stillicidiorum 25 ‘4’
C. stillicidiorum 26 ‘3’
C. stillicidiorum 27 ‘3’
C. stillicidiorum 28 ‘4’
C. stillicidiorum 29 ‘4’
C. stillicidiorum 30 ‘4’
C. stillicidiorum 31 ‘4’
C. stillicidiorum 32 ‘4’
C. stillicidiorum 33 ‘4’
C. stillicidiorum 34 ‘5’
C. stillicidiorum 35 ‘5’
C. stillicidiorum 36 ‘6’
C. stillicidiorum 37 ‘6’
C. stillicidiorum 38 ‘6’
C. stillicidiorum 39 ‘6’
C. stillicidiorum 40 ‘6’
C. stillicidiorum 41 ‘5’
C. stillicidiorum 42 ‘5’
C. stillicidiorum 43 ‘5’
C. stillicidiorum 44 ‘5’
C. stillicidiorum 45 ‘5’
C. stillicidiorum 46 ‘5’
C. stillicidiorum 47 ‘5’
C. stillicidiorum 48 ‘5’
*
Specimen
Bulgaria, Vondrák JV1992 (CBFS)
Russia, Kondratyuk 20709 (KW)
Austria, Hafellner JH42134 (GZU)
Austria, Vondrák JV3821 (CBFS)
Austria, Kocourková 900101 (PRM)
Austria, Hafellner & Muggia JH67073 (GZU)
Sweden, Arup L04172 (LD)
Alaska, Søchting 7440 (C)
Svalbard (Norway), Košnar 667 (hb. Šoun)
Greenland (Denmark), Hansen 7525 (C)
Greenland (Denmark), Hansen 04074 (C)
USA, Montana, Spribille 14195 (hb. Spribille)
Canada, British Columbia, Spribille & Goward 15291
(hb. Spribille)
Norway, Košnar 669 (hb. Šoun)
Norway, Košnar 668 (hb. Šoun)
Czech Republic, Vondrák JV6330 (CBFS)
Czech Republic, Vondrák & Šoun JV2911 (CBFS)
Czech Republic, Vondrák & Šoun JV2910 (CBFS)
Ukraine, Khodosovtsev 2937 (KHER)
Kazakhstan, Khodosovtsev & Vondrák JV7644 (CBFS)
Austria, Vondrák JV2974 (CBFS)
Romania, Vondrák JV3617 (CBFS)
Ukraine, Khodosovtsev 2938 (KHER)
Czech Republic, Vondrák JV4824 (CBFS)
Czech Republic, Kocourková & Šafránek 908963 (PRM)
Ukraine, Nadyeina JV7620 (CBFS)
Hungary, Vondrák JV4867 (CBFS)
Hungary, Vondrák & Šoun JV4388 (CBFS)
Spain, Vondrák JV2612 (CBFS)
Finland, Palice 11679 (hb. Palice)
Turkey, Šoun 246 (hb. Šoun)
Spain, Søchting 9685 (C)
Uzbekistan, Mishutin 2940 (KHER)
USA, Montana, Spribille & Goward 10676 (hb. Spribille)
Spain, Vondrák JV2690 (CBFS)
Turkey, Šoun 247 (hb. Šoun)
GenBank accession
number
HM538464
EU681283
HM538569
HM538570
HM538573
HM538572
HM538571
HM538568
HM538575
HM538574
HM538550
HM538556
HM538557
HM538576
HM538577
HM538551
HM538553
HM538552
HM538554
HM538549
HM538555
HM538559
HM538558
HM538456
HM538460
HM538459
HM538458
HM538457
HM538560
HM538561
HM538562
HM538564
HM538563
HM538565
HM538566
HM538567
Sequences have an intron 218 bases long in the 18S region
Sequence alignment and phylogenetic analyses
One hundred and twenty two newly obtained ITS
sequences were included in the phylogenetic analysis
along with seventeen sequences of the Caloplaca cerina
group available from GenBank (Table 2). Sequences
of Caloplaca aurantia (AY233219) and C. flavescens
(EU639601) from GenBank were selected as an outgroup. Sequences were aligned using MAFFT 6 (on-line
version in the Q-INS-i mode; see Katoh et al. 2002) and
manually cut to eliminate the unalignable ends, the
intron and ambiguously aligned regions of ITS1 and
ITS2. The final alignment included 596 positions.
Bayesian phylogenetic analysis was carried out using the
programme MrBayes 3.1.1 (Ronquist & Huelsenbeck
2003). The General Time Reversible substitution
model (Rodriguez et al. 1990) with estimation of invariant sites and assuming a gamma distribution with four
categories (GTR+I+G) was used for likelihood calculations. The optimal nucleotide substitution model was
found with the program MrModeltest v2.3 (Nylander
2004) using the Akaike Information Criterion and the
hierarchical likelihood ratio test (Posada & Crandall
1998). The MCMC analysis was run for ten million
generations, performed in two runs, each with four
chains starting from a random tree and using the
default temperature of 0·2. Every 100th tree was sampled, and the first 30% of the trees were discarded as
burn-in.
2011
The Caloplaca cerina group in Europe—Šoun et al.
119
T 3. Anthraquinone contents (% concentrations) of selected species of the Caloplaca cerina group
C. cerina A (CBFS JV2548)
C. cerina A (CBFS JV3776)
C. cerina A (hb. Palice 11906)
C. cerina B (LD L02355)
C. cerina C (hb. Šoun 253)
C. cerina D (hb. Šoun 250)
C. chlorina (CBFS JV1879)
C. isidiigera (hb. v. d. Boom 10756)
C. monacensis (CBFS JV548)
C. pinicola (GZU 80302)
C. aff. pinicola (hb. Spribille 4731)
C. stillicidiorum 2 (hb. Šoun 667)
C. stillicidiorum 6 (hb. Šoun 61)
C. turkuensis (CBFS JV6868)
C. ulmorum (GZU 90256)
emodin
fallacinal
parietin
parietinic acid
4
1
4
5
2
2
4
3
1
4
2
2
2
5
2
4
4
4
3
5
3
9
4
3
3
2
5
4
4
3
89
91
90
91
86
92
85
91
95
92
94
91
83
89
93
1
3
1
1
2
1
0
1
1
1
1
2
1
2
1
Chemistry
Lichen substances in the apothecia were extracted in
150 l of acetone at room temperature. The extract was
subjected to high-performance liquid chromatographic
analysis (HPLC). Reverse phase column (C18, 5 m,
Lichrocart 250-4) was eluted with MeOH/30%MeOH+
1%H3PO4 for 77 min and the absorbance at 270 nm was
recorded (for details see Søchting 1997). The compounds were determined on the basis of their retention
times and absorption spectra. Acetone insoluble pigments were examined according to Meyer & Printzen
(2000).
Results
Phylogeny
The final alignment of 596 nucleotides had
134 variable positions. The following species
had an insertion 218 bases long in the 18S
region: C. stillicidiorum (only ecotype 2), C.
pinicola, C. aff. pinicola, C. monacensis (except
van den Boom 38821 and CBFS JV3236),
C. ulmorum, C. thracopontica, C. aff. thracopontica, C. sterilis, C. hanneshertelii and C. sp.
(Frödén 1246, LD). The insertion was not
included in the analysis.
The phylogenetic tree, based on the
Bayesian inference analysis is presented in
Fig. 1 along with the posterior probabilities
for the nodes. The species C. isidiigera,
C. sterilis, C. subalpina, C. turkuensis and
teloschistin unknown
2
1
0
0
3
1
2
1
0
0
1
0
10
0
1
0
0
1
0
2
1
0
0
0
0
0
0
0
0
0
C. ulmorum form well-supported clades
(PP > 0·95) in the cladogram. The species
C. chlorina, C. monacensis and C. thracopontica are not so well supported. Caloplaca
cerina s. lat. is a polyphyletic taxon with six
lineages, possibly cryptic or semi-cryptic
species. Caloplaca stillicidiorum s. lat. is a
monophyletic taxon (PP = 0·92) forming
the crown group of the tree. This clade is
internally diverged into three major lineages
(ecotype 1, ecotype 2, ecotype 3–4, ecotype
5–6), possibly representing several semicryptic species. The only available sequence
of C. pinicola has a sister relationship to
three similar sequences of North American
sorediate C. cerina-like specimens which
probably represent an unknown species
(see the note under C. pinicola). The only
sequence of C. aff. thracopontica from the
Mediterranean region is close to the C.
thracopontica clade but possibly not conspecific with it. One available sequence of
the Australian C. hanneshertelii and one
sequence of an undescribed species from
Tasmania confirmed their affinities to the
C. cerina group.
Chemistry
The anthraquinone content of all species
analyzed from the C. cerina group is uniform
120
THE LICHENOLOGIST
C. aurantia
C. flavescens
0.66
C. cerina 1
C. cerina 2
C. cerina 3
C. cerina 4
C. cerina 5
0.67
C. cerina 6
C. cerina 7
0.96
Vol. 43
A
C. sp. (Tasmania)
C. subalpina 1
C. subalpina 2
C. subalpina 3
1.00
C. cerina 8
C. cerina 9
B
C. cerina 10
C. cerina 11
C. cerina 12
C
C. cerina 13
C. turkuensis 1
C. turkuensis 2
1.00
C. turkuensis 3
C. turkuensis 4
0.67
C. turkuensis 5
C. turkuensis 6
C. hanneshertelii
C. isidiigera 1
C. isidiigera 2
C. isidiigera 3
C. isidiigera 4
1.00
C. isidiigera 5
0.96
C. isidiigera 6
C. isidiigera 7
C. isidiigera 8
1.00
0.61
C. isidiigera 9
C. isidiigera 10
C. ulmorum 1
0.99
1.00
C. ulmorum 2
C. ulmorum 3
0.97
0.66
C. monacensis 1
C. monacensis 2
C. monacensis 3
0.57
C. monacensis 4
C. monacensis 5
0.55
0.83
C. monacensis 6
C. monacensis 7
0.86
1.00
C. monacensis 8
C. monacensis 9
C. monacensis 10
0.95
C. monacensis 11
C. monacensis 12
0.90
C. monacensis 13
0.99
C. monacensis 14
0.96
C. monacensis 15
C. monacensis 16
C. aff. thracopontica
1.00
0.89
1.00
0.90
0.92
C. sterilis 1
1.00
C. sterilis 2
C. sterilis 3
C. sterilis 4
C. thracopontica 1
C. thracopontica 2
0.88
C. thracopontica 3
0.68
C. thracopontica 4
1.00
C. thracopontica 5
C. thracopontica 6
C. thracopontica 7
C. chlorina 1
0.95
C. chlorina 2
1.00
C. chlorina 3
0.52
C. chlorina 4
C. chlorina 5
C. chlorina 6
1.00
C. chlorina 7
C. chlorina 8
0.87
C. chlorina 9
C. chlorina 10
0.97
C. chlorina 11
C. chlorina 12
C. chlorina 13
C.
pinicola
0.96
C. aff. pinicola 1
1.00
C. aff. pinicola 2
C. aff. pinicola 3
C. cerina 14
0.99
0.60
0.81
D1
C. cerina 15
C. cerina 16
0.99
C. cerina 17
0.93
C. cerina 18 D2
C. cerina 19
0.86
C. cerina 20
0.98
0.68
C. cerina 21
D3
0.89
C. cerina 22
C. stillicidiorum 1
1.00
C. stillicidiorum 2
C. stillicidiorum 3
C. stillicidiorum 4
0.81
1.00
1.00
C. stillicidiorum 5
C. stillicidiorum 6
C. stillicidiorum 7
1
0.73
C. stillicidiorum 8
C. stillicidiorum 9
1.00
C. stillicidiorum 10
0.89
C.
stillicidiorum
11
0.92
C. stillicidiorum 12
C. stillicidiorum 13
C. stillicidiorum 14
0.76
C. stillicidiorum 15
1.00
C. stillicidiorum 16
2
0.85
C. stillicidiorum 17
C. stillicidiorum 18
C. stillicidiorum 19
C. stillicidiorum 20
C. stillicidiorum 21
3
C. stillicidiorum 22
0.62
0.64
C. stillicidiorum 23
1.00
C. stillicidiorum 24
4
0.99
C. stillicidiorum 25
1.00
C. stillicidiorum 26
C. stillicidiorum 27 3
C. stillicidiorum 28
C. stillicidiorum 29
1.00
0.55
C. stillicidiorum 30
C. stillicidiorum 31 4
0.78
C. stillicidiorum 32
C. stillicidiorum 33
C. stillicidiorum 34
5
C. stillicidiorum 35
C. stillicidiorum 36
C. stillicidiorum 37
1.00
1.00
C. stillicidiorum 38
6
C. stillicidiorum 39
C. stillicidiorum 40
C. stillicidiorum 41
C.
stillicidiorum
42
0.81
C. stillicidiorum 43
0.96
C. stillicidiorum 44
1.00
C. stillicidiorum 45
C. stillicidiorum 46
0.91
0.90
C. stillicidiorum 47
C. stillicidiorum 48
0.99
0.93
5
0.1
F. 1. Bayesian consensus phylogeny of the Caloplaca cerina group inferred from nrDNA ITS sequences. Node
support values are Bayesian posterior probabilities.
2011
The Caloplaca cerina group in Europe—Šoun et al.
(Table 3) and corresponds with chemosyndrome A (Søchting 1997). Sedifolia-grey, a
pigment insoluble in acetone, is present in
the cortex of the thallus, the thalline margin
121
and in the vegetative diaspores, except the
soredia of C. sterilis, of all studied species
from the group (C+, K+, N+ pinkish violet/
violet/sordid violet in section).
Key to fertile species known to belong to the Caloplaca cerina group. Sterile
specimens can be confused with various species outside the group or even outside
the genus Caloplaca
1
Ascospore septum relatively thin, 1·5–4·0 m; thallus dark grey; soredia often
present; North America . . . . . . . . . . . . . . . . . . . . . . . . C. pinicola
Ascospore septum wider, 4·0–7·0 m . . . . . . . . . . . . . . . . . . . . . . . . 2
2(1)
Thallus without any vegetative diaspores . . . . . . . . . . . . . . . . . . . . . . 3
Thallus with vegetative diaspores (lobules, isidia, granules, soredia/blastidia). . . 4
3(2)
On trees and shrubs . . . . . . . . . . . . . . . . . . . . . . . . C. cerina s. lat.
On mosses, plant debris, wood, bark of exposed roots, dwarf-bushes and rarely on
rocks . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stillicidiorum s. lat.
4(2)
Thallus with lobules, isidia or granules (usually > 50 m diam.) . . . . . . . . . 5
Thallus with soredia/blastidia (usually up to 50 m diam.) . . . . . . . . . . . . 9
5(4)
Thallus surface with pustules and lobules, 100–400 m wide or rarely without any
vegetative diaspores; saxicolous, maritime . . . . . . . . . . . C. thracopontica
Thallus with isidia or granules. . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6(5)
Thallus with isidia; apothecia absent or present . . . . . . . . . . . . . . . . . . 7
Thallus with granules on margins of areoles or completely granulose; granules (53–)
96 ± 27 (–163) m diam.; usually richly fertile . . . . . . . . . . . . . . . . . 8
7(6)
Thallus areolate covered by small globose to shortly vertically elongated isidia, (25–)
66 ± 20 (–140) m wide . . . . . . . . . . . . . . . . . . . . . . C. isidiigera
Thallus with branched coralloid isidia and erect lobules . . . . .C. squamuloisidiata
(species with unsettled phylogenetic position)
8(6)
Thallus areolate with marginal granules, rarely predominantly granular; corticolous;
North America. . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ulmorum
Thallus nearly to completely granulose; corticolous, rarely saxicolous; Europe,
Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. monacensis
9(4)
Soralia crater-like, Australia . . . . . . . . . . . . . . . . . . . C. hanneshertelii
Soralia different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10(9)
Areoles at thallus margin forming squamules, (0·16–) 0·58 ± 0·32 (–2·04) mm wide;
thallus, at least in marginal parts, white-pruinose . . . . . . . . . C. subalpina
Thallus without marginal squamules . . . . . . . . . . . . . . . . . . . . . . . 11
11(10) Thallus with white-pruinose delicate areoles, (0·05–) 0·16 ± 66 (–0·30) mm wide,
soralia (sordid) grey-green to pale green soralia . . . . . . . . . . . . C. sterilis
Thallus without pruina; soralia darker, usually with grey or blue tinge . . . . . 12
122
THE LICHENOLOGIST
Vol. 43
12(11) Areoles usually relatively large, up to 1·2 mm wide; soredia/blastidia (15–) 33 ± 7
(–53) m diam.; apothecia common, apothecial margin not sorediate; saxicolous
or on nutrient-rich tree bases . . . . . . . . . . . . . . . . . . . . . C. chlorina
Thallus with very small areoles up to 0·2 mm wide; areoles usually very soon entirely
sorediate; soredia smaller (15–) 22 ± 3 (–30) m diam.; apothecial margin often
sorediate, at least in part, however some specimens completely without sorediate
margin; corticolous, lignicolous . . . . . . . . . . . . . . . . . . .C. turkuensis
(morphologically resembles another phylogenetically distant undescribed Caloplaca species, differing in
soralia and apothecia with an endophloeic origin, not developing from areoles, the true exciple usually
more apparent and the thalline margin not so stout)
The Species
Caloplaca cerina s. lat.
Including Caloplaca cerina (Hedw.) Th. Fr. s. str., C.
gilva (Hoffm.) Zahlbr., C. cerina var. azaleae (Vain.)
Zahlbr., C. cerina var. circumponens (Nyl.) Boist., C.
cerina var. subfuscescens (Nyl.) Blomb., C. cerina f.
chrysaspis (Ach.) Th. Fr., C. cerina f. corni Britzelm.
This is a polyphyletic taxon split into six
different clades scattered within the cladogram, comprising corticolous specimens
lacking vegetative diaspores. Three of the
clades, D1, D2 and D3, are very closely
related and seem to have similar distribution and ecology; they are here treated together (group D). We keep all the lineages
unnamed, without nomenclatural status,
because we failed to find sufficient morphological characters to separate them. Brief
summaries of the characteristics of each of
the molecularly based groups are presented
here:
A: on bark of Cupressus, Olea, Pistacia,
and Platanus in the Mediterranean region
(Greece, Spain, Turkey). Thallus as in clade
B but tends to be darker; apothecia yellow to
orange-yellow, strongly pruinose on disc and
inner part of margin, to 1·1 mm diam.;
ascospores (12–) 13·8 ± 1·1 (–16) × (5–)
6·4 ± 0·7 (–8) m; septum (5–) 5·9 ± 0·7
(–7) m wide (n = 60).
B: on bark of deciduous trees in northern
Iran. Thallus crustose, thick, pale grey to
beige, sometimes whitish; apothecia orangeyellow to deep orange, disc sometimes
slightly pruinose when young, to 0·9 mm in
diam.; ascospores (14–) 14·8 ± 0·8 (–17) ×
(7–) 8·3 ± 1·1 (–11) m; septum (4–)
5 ± 0·5 (–6) m wide (n = 20).
C: on bark of Salix in Scandinavia
(Norway, Sweden) and on Fraxinus in
Hyrcanian forest in Iran. Thallus and apothecia similar to the type D but smaller; apothecia orange-yellow, not pruinose, to 0·8 mm
diam.; ascospores (10–) 12·6 ± 2 (–17) ×
(5–) 6·8 ± 1 (–9) m; septum (3–) 4·1 ± 1
(–6) m wide (n = 30).
D1, D2 and D3: on bark of Acer, Juglans,
Populus, Prunus, Sambucus throughout
Europe (Austria, Czech Republic, Finland,
Greece, Norway, Slovakia, Spain). Thallus
crustose, pale to dark grey, sometimes
warted, thick or thin; apothecia orangeyellow to orange, rarely pruinose, to 1·6 mm
diam.; ascospores (11–) 14·6 ± 1·7 (–20) ×
(6–) 7·9 ± 0·9 (–10) m; septum (5–) 6·7 ±
1·2 (–10) m wide (n = 90).
The holotype of C. cerina (Hedw.) Th. Fr.
is the illustration in Hedwig (1788: 62, tab.
21, f. B). Wetmore (2007a) designated an
epitype (MIN!) and several isoepitypes
(ASU, COLO, MIN, US) using the exsiccate Lichenotheca Fenn. 116 (Finland, Ostrobottnia Borealis: Simo, insula Kumurainen,
leg. Räsänen). The type illustration is difficult to assign to any of the clades described
above. The epitype probably belongs to clade
C because of its relatively small apothecia
(to 1 mm diam.) and small spores (11–13 ×
7–8 m, septum 5–5·6 m). However, a
placement in the other Scandinavian clades,
D1 or D3, is also possible.
Caloplaca chlorina (Flot.) H. Olivier
Mém. Soc. natl. Sci. nat. et math. Cherbourg 37: 122
(1909).—Zeora cerina * [var.] chlorina Flot., Jahresber.
Schlesisch. Gesellsch. für vaterl. Kultur 27: 126 (1849);
type: Callopisma cerinum a** chlorinum, Körb. Lichenes
Selecti Germanici Exsiccati no. 128 (1857), ad saxa
2011
The Caloplaca cerina group in Europe—Šoun et al.
123
F. 2. Caloplaca species. A, C. chlorina (CBFS JV2055); B, C. isidiigera (Šoun 283); C, C. monacensis (v. d. Boom
38821); D, C. monacensis Arnold Lich. Monacenses exs. 1896 no. 422, note apothecial discs without normal
yellow-orange colour (BP—isotype); E. & F, C. turkuensis (CBFS JV2921). Scales: A–F = 1 mm.
granitica circa “Drachenburg” prope Hirschbergam
Silesiae, Körber [O—neotype designated by Wetmore
(1997), L, M!, MIN, S!—isoneotypes].
Caloplaca chlorina var. cyanopolia (Nyl.) H. Olivier,
Mém. Soc. natl. Sci. nat. math. Cherbourg 37: 123
(1909).—Lecanora cerina f. cyanopolia Nyl., Notiser ur
Sällsk. pro Fauna et Flora Fennica Förhandlingar 8: 128
(1866); type: Russia: Republic of Karelia, Sortavala,
1863, Kullhem [H-NYL 29770!—lectotype designated
by Kärnefelt (1990)].
Caloplaca chlorina var. paradoxa (Vain.) H. Olivier,
Mém. Soc. natl. Sci. nat. math. Cherbourg 37: 123
(1909).—Lecanora cerina var. cyanopolia *[f.] paradoxa
Vain., Meddel. Societas pro Fauna et Flora Fennica 3: 55
(1878); type: Karelia australis, Wiipuri, Monrepos
[Russia, Leningrad Oblast, Vyborg], 1875, Lang
(TUR-V 07552!).
(Fig. 2A)
124
THE LICHENOLOGIST
Thallus areolate, up to several cm wide,
non-pruinose, up to 0·27 (–0·53) mm thick,
always sorediate/blastidiate; sometimes also
with tiny isidiate outgrowths especially at
margins of older areoles; areoles flat to convex, up to 1·2 mm wide; colour variable,
sordid white, pale grey, brownish grey, dark
grey to almost black, usually with bluish
tinge; soralia and blastidia pale to dark grey,
with bluish tinge, rarely greenish grey, arising
from margins of areoles, sometimes spreading over the whole areole; soredia/blastidia
(15–) 33 ± 7 (–53) m diam. (n = 100);
soredia sometimes forming consoredia; prothallus sometimes present, thin and black;
cortex up to 10 (–30) m thick, paraplectenchymatous.
Apothecia lecanorine, abundant to absent,
up to 1·5 mm diam., sessile, non-pruinose;
disc flat to slightly convex, pale to dark
orange; true exciple indistinct, very thin, up
to 15 m thick, prosoplectenchymatous;
thalline exciple of thallus colour, raised above
disc when young, somewhat reduced in old
apothecia, 60–90 m thick; epihymenium
orange, with inspersed granules; hymenium
50–85 m thick, hyaline; hypothecium (55–)
80–110 m thick, hyaline; paraphyses simple
to branched, 2 m broad with upper cells
wider, 3–5 m thick; asci 8-spored, 35–
59 × 10–18 m (n = 50); ascospores polarilocular, ellipsoid, (10·0–) 12·4 ± 0·9
(–15·0) × (5·0–) 6·6 ± 0·6 (–8·0) m; septum (4·0–) 4·7 ± 0·5 (–6·0) m wide
(n = 100); ratio of septum width/spore
length (0·3–) 0·4 ± 0·04 (–0·5).
Pycnidia rare to scattered, immersed,
distinct by elevated darker ostiole; conidia
bacilliform, 3–4 × 1 m.
Habitat and distribution. This species grows
from lowlands to mountains on basic or
nutrient enriched siliceous (e.g. gneiss, slate,
granite) and limestone rocks both in natural
and anthropogenic habitats, especially on old
stone walls, pebbles in railroads, bricks,
roofing-tiles and mortar. It is occasionally
found on bark of broad-leaved trees (e.g.
Acer, Fraxinus, Juglans, Ulmus) on trunk
bases. It seems to prefer humid or shaded
sites and substrata near the ground. Probably
Vol. 43
it is quite frequent throughout Europe but
often overlooked or confused with similar
taxa. There are confirmed records from
Austria, Belgium, Bulgaria, Czech Republic,
Denmark, Germany, Ireland, Netherlands,
Norway, Portugal, Romania, Russia,
Slovakia, Sweden, Ukraine and the United
Kingdom. The world distribution (see
Fletcher & Laundon 2009) needs revision
but three records from North America are
confirmed.
Remarks. The name C. chlorina has often
been used in a broad sense for saxicolous and
corticolous specimens of the C. cerina group
with any vegetative diaspores. Wetmore
(1997) selected a neotype for C. chlorina
and stated that the isidiate C. isidiigera is a
synonym. However, according to our results,
C. chlorina s. str. is always characterized by
the presence of soredia/blastidia. Only some
morphotypes of C. chlorina, usually overgrown or grazing affected thalli, possessing
various isidia-like secondary outgrowths and
consoredia, can resemble C. isidiigera. Sterile
specimens can be confused with some unrelated species, for example C. soralifera
Vondrák & Hrouzek.
Selected specimens examined. Austria: Lower Austria:
Wolfsthal, castle ruins Pottenburg, Vondrák JV1879
(CBFS). Carinthia: Gailtaler Alpen, Hermagor,
Weibriach, van den Boom 16234 (hb. v. d. Boom).
Burgenland: Bernsteiner Gebirge, Bernstein, Hafellner &
Maurer JH31242 (GZU).—Belgium: Luxembourg:
Martelange, Witry, van den Boom & van den Boom
30644 (hb. v. d. Boom).—Bulgaria: Haskovo: Eastern
Rodopi Mts, Malko Gradishte, Vondrák JV2226
(CBFS); Madzharovo, Vondrák JV2055 (CBFS).—
Canada: British Columbia: Wells Gray Educ. and Res.
Centre, Søchting 7086 (C). Newfoundland: Waghorne
(BM).—Czech Republic: Central Bohemian Reg.:
Rakovnı́k, castle ruins Týřov, Vondrák JV2982, JV2764
(CBFS). South Bohemian Reg.: Pı́sek, castle Zvı́kov,
Vondrák JV3034, (CBFS). Zlín Reg.: Kroměřı́ž,
castle Buchlov, Vondrák JV1294 (CBFS). Liberec Reg.:
Jizerské hory Mts, Jizerka, Kocourková & Kocourek
907659 (PRM). Karlovy Vary Reg.: Karlovy Vary,
Andělská Hora, Vondrák JV1103 (CBFS). Plzeň Reg.:
Konstantinovy Lázně, castle ruins Krası́kov, Vondrák
JV2294 (CBFS). Vysočina Reg.: Nové Město na Moravě,
Kut’ák 762037 (PRM). Moravian-Silesian Reg.: Hrubý
Jesenı́k Mts, Mt Vysoká hole, Vondrák JV1932 (CBFS).
South Moravian Reg.: Kyjov, Milotice, Vondrák JV1373
(CBFS).—Denmark: Bornholm: Hammershus, Larsen
& Vondrák JV4714 (CBFS).—Germany: Baden-
2011
The Caloplaca cerina group in Europe—Šoun et al.
Württemberg: Heidelberg (M).—Ireland: Connemara:
Lough Feagh and Lough Muck, Larbalestier 23 (BM).—
Netherlands: North Holland: Naarden-Vesting, van
den Boom 15417 (hb. v. d. Boom). North Brabant:
Werkendam, Woudrichem, van den Boom 12545 (hb.
v. d. Boom).—Norway: Aust-Agder: Grimstad, Tønsberg
13551 (BM).—Portugal: Trás-os-Montes: Bragança,
Zeive, van den Boom 19357 (hb. v. d. Boom).—
Romania: Arad: Arad, Nǎdlac, Vondrák JV2222
(CBFS).—Slovakia: Trenčín Reg.: Považský Inovec
Mts, Tematı́n, Halda & Palice 10186 (hb. Palice).
Bratislava Reg.: Bratislava, Mt Devı́nská kobyla,
Vondrák JV1822 (CBFS).—Sweden: Värmland:
Frykerud, Arup L03566 (LD). Skåne: Maltesholm,
Hessikahuset, Arup L08033 (LD).—Ukraine: Crimea:
Karadag Mts, Mt Svyataya, Vondrák 290 (hb. Šoun).—
Great Britain: Scotland: Dumfries, Drumlanrig Castle,
James (BM); Banff, Mountblairy, James (BM); Mid
Ebudes, Ballyhaugh, James (BM). Wales: Monmouthshire, Abergavenny, James (BM).—USA: Montana:
Lincoln Co., W of Libby, Spribille 11895 (hb. Spribille).
Exsiccata examined. Finland: Finland Proper Reg.:
Kakskerta, Lich. Fenn. Exs. no. 234 [as Placodium gilvum
var. chlorina].—Russia: Karelia: Kurkiyeki, Lich. Fenn.
Exs. no. 85 [as Placodium gilvum var. chlorina] (BM).—
Sweden: Närke: Mellösa, Arnold: Lich. Exs. no. 1550a
[as Callopisma cerinum f. chlorinum] (BM).
Caloplaca hanneshertelii S.Y. Kondr. &
Kärnefelt
Bibliotheca Lichenologica 88: 262 (2004); type: Australia,
Victoria, NW of Swan Hill, 22 km W of Nyah West,
on shrubs along the road, c. 50 m alt., 35°10#61$S,
143°09#14$E, growing together with Xanthoria filsonii,
15 January 1999, Kärnefelt 992101 (LD—holotype).
This Australian corticolous species is characterized by crater-like soralia with darkbluish soredia erupting from pustule-like
outgrowths at whitish grey areoles. Apothecia are small, 0·2–0·5 mm diam. Caloplaca
ahtii Søchting, C. obscurella (J. Lahm) Th. Fr.
and C. ulcerosa Coppins & P. James have
similar type of soralia but their apothecia
differ. For detailed description of this species
see Kärnefelt & Kondratyuk (2004).
Caloplaca isidiigera Vězda
Folia Geobot. Phytotax. Bohemoslov. 13: 417 (1978);
type: Caloplaca isidiigera, Vězda Lich. Sel. Exs. no. 1494,
Czechoslovakia, Eu-Carpaticum, Tatra Minor, in
monte Velký bok, c. 1550 m s. m., ad saxa calcarea,
September 1974, Vězda (PRA-V—holotype; PRM!,
BM!—isotypes).
(Fig. 2B)
125
Thallus areolate, up to several cm wide,
non-pruinose, up to 0·4 (–0·8) mm thick,
isidiate; areoles convex when very young, later
flat, up to 2·1 mm wide; colour varies from
sordid pale grey, dark grey to black, usually
with bluish tinge; lower part of very thick
areoles often stained purple-black (in section); isidia initially arising mainly from margins of areoles, but soon covering whole
surface of areoles, globose to shortly elongated or flattened, concolorous with the
areoles; (25–) 66 ± 20 (–140) m diam.
(n = 82); prothallus usually present, thin and
black; cortex up to 20 m thick, paraplectenchymatous.
Apothecia lecanorine, abundant to absent,
up to 1·3 mm diam., sessile, non-pruinose;
disc flat to slightly convex, pale to dark
orange; true exciple indistinct, very thin, up to
10 m thick, prosoplectenchymatous; thalline exciple of thallus colour, raised above disc
when young, somewhat reduced in old
apothecia, sometimes crenulate to isidiate,
65–110 m thick; epihymenium orange, with
inspersed granules; hymenium 55–85 m
thick, hyaline; hypothecium 60–110 m thick,
hyaline; paraphyses simple to branched, 2 m
broad with upper cells wider, 3–5 m thick;
asci 8-spored, 38–55 × 9–15 m (n = 24);
ascospores polarilocular, ellipsoid, (10·0–)
12·4 ± 1·4 (–15·0) × (5·0–) 6·5 ± 0·9 (–8·0)
m; septum (3·0–) 4·6 ± 0·7 (–6·0) m wide
(n = 48); ratio of septum width/spore length
(0·3–) 0·4 ± 0·05 (–0·5).
Pycnidia rarely seen, immersed; conidia
bacilliform, 3 × 1 m.
Habitat and distribution. The species seems
to grow only in subalpine and alpine habitats
on natural limestone, dolomite or calciferous
slate and base-rich sandstone rocks. Confirmed records are from the Alps (Austria,
Germany, Italy and Switzerland), Carpathians (Slovakia, Ukraine) and Scandinavia
(Sweden). One record is confirmed from
North America.
Remarks. This species is characterized by
the presence of isidia and the type specimens
are clearly isidiate. However, Wetmore
(1996) considered these structures seen in
126
THE LICHENOLOGIST
the isotypes as lobules and van den Boom &
Rico (2006) regarded them as granules or
microsquamules. This has caused many
misinterpretations and the name has, for
example, been applied to the corticolous C.
monacensis. Sterile specimens might be confused with unrelated C. xerica Poelt & Vězda,
but it differs in growing on base-rich siliceous
rocks in xerothermic habitats.
Nomenclatural note. Caloplaca isidiigera was
synonymized by Wetmore (1997) with C.
chlorina, and thereafter not accepted as a
separate species. This study clearly shows
that both taxa are separate species (for more
details see the note under C. chlorina).
Selected specimens examined. Austria: Styria:
Ennstaler Alpen, Reichenstein-Massiv, Hafellner,
Muggia & Hafellner JH68548 (GZU); Schladminger
Tauern, Ursprungalm, Türk RT32349 (GZU);
Gurktaler Alpen, Turrach, Hafellner JH61513 (GZU);
Steirisches Randgebirge, Grazer Bergland, Mixnitz,
Hafellner JH64492 (GZU); Nördliche Kalkalpen,
Hochschwab-Gruppe, Trenchtling, Hafellner &
Hafellner JH62455 (GZU); Mürzsteger Alpen,
Schneealpe, Mayrhofer & Bilovitz 17752 (GZU);
Wölzer Tauern, Lachtal, Mayrhofer & Sterner 13973
(GZU); Stubalpe, Wölkerkogel, Wilfling & Unger
AW2315 (GZU); Niedere Tauern, Gumpeneck,
Hafellner & Wilfling AW1728 (GZU); Eisenerzer Alpen,
Reiting-Massiv, Hafellner & Hafellner JH43577 (GZU).
Carinthia: Karnische Alpen, Schloßhüttensattel, Poelt
IN52-84 (GZU). Tyrol: N. P. Hohe Tauern, Kals,
Hafellner JH47074 (GZU).—Germany: Bavaria:
Berchtesgaden N. P., Türk & Wunder (M).—Italy:
Trentino-Alto Adige: Merano, NP Stilfser Joch, van den
Boom 10756 (hb. v. d. Boom).—Slovakia: Žilina Reg.:
Nı́zke Tatry Mts, Mt Vel’ký bok, Šoun 283 (hb.
Šoun).—Sweden: Lule Lappmark: Padjelanta national
park, Allak, Arup L04227 (LD). Torne Lappmark:
Torneträsk, Bessisjohka, Arup L04228 (LD). Pite
Lappmark: Arjeplog par., Mt Skärrim, Arup L03570
(LD). Härjedalen: NE of Ramundberget, Mt StorMittåkläppen, Arup L02352 (LD).—Switzerland:
Berner Oberland: Brienz, Chüemad, van den Boom 17968
(hb. v. d. Boom). Valais: Val de Moiry, Fêta d’Août de
Chateaupré, van den Boom 10108 (hb. v. d. Boom).—
Ukraine: Zakarpatska oblast: Svidovets Mts, Vondrák
JV6073 (CBFS).—USA: Montana: Lincoln Co., Whitefish Range, Mt Marston, Spribille 12617 (hb. Spribille).
Exsiccata examined. Ukraine: Zakarpatska oblast:
Svidovets Mts, Vondrák: Sel. Exs. of Caloplaca no. 42
(CBFS).
Caloplaca monacensis (Leder.) Lettau
Hedwigia 52: 240 (1912).—Pyrenodesmia monacensis
Leder., Bericht. Bayr. Botan. Gesellsch. 4: 26 (1896);
Vol. 43
type: Pyrenodesmia monacensis, Arnold Lich. Monacenses
Exs. 1896 no. 422, an alten Strassenpappeln nicht weit
vom Warthof bei Giesing, München, März 1896, Lederer
[M (M-0023624)!—lectotype designated here, BP!,
BM!, M (M-0053460)!—isolectotypes].
(Figs 2C & D)
Thallus predominantly to entirely granular,
scarcely with distinct areoles, especially
around apothecia, scattered to continuous,
thin to thick, non-pruinose, (sordid) pale to
dark grey; granules (53–) 96 ± 27 (–163) m
diam. (n = 99). Sometimes very minutely
tomentose with inconspicuous hairs 4–10 ×
2–3 m formed by projecting hyphae (visible
in section), particulary on granules; prothallus
not seen.
Apothecia lecanorine, abundant to rare, up
to 1·2 mm diam., sessile, distinct, sometimes
pruinose; disc flat to slightly convex, yelloworange to orange, sometimes pruinose,
especially when young; true exciple indistinct,
very thin, up to 20 m thick, prosoplectenchymatous; thalline exciple of thallus colour
and often whitish pruinose, raised above the
disc when young, somewhat reduced in old
apothecia, 55–110 m thick; epihymenium
orange, with inspersed granules; hymenium
(65–) 85–102 m thick, hyaline; hypothecium
(45–) 75–90 m thick, hyaline; paraphyses
simple to branched, 2 m broad with upper
cells wider, 4–6 m diam.; asci 8-spored,
40–67 × 10–22 m (n = 48); ascospores
polarilocular, ellipsoid, (11·0–) 13·4 ± 1·3
(–16.0) × (5·0–) 6·5 ± 0·7 (–8·0) m, septum (3·0–) 4·9 ± 0·7 (–7·0) m wide
(n = 100), ratio of septum width/spore
length (0·2–) 0·4 ± 0·05 (–0·5).
Pycnidia rare to scattered, immersed,
made distinct by elevated darker ostiole;
conidia bacilliform, 3–4 × 1 m.
Habitat and distribution. This species grows
on bark of wayside trees, solitary trees in
pastures and in other types of agricultural
landscapes and in open deciduous or mixed
forests. It has been recorded on a variety
of broad-leaved trees such as Acer, Fagus,
Fraxinus, Juglans, Ostrya, Pistacia, Populus,
Pyrus, Quercus, Salix, Tilia, Ulmus, but it is
also rarely found on bark of shrubs and rotten
2011
The Caloplaca cerina group in Europe—Šoun et al.
wooden fences. Once it was found growing
on shaded cherty limestone in Bükk Mts,
Hungary (Vondrák et al. 2009b). Probably it
is widely distributed from the lowlands to
medium altitudes in the mountains throughout temperate and Mediterranean Europe
and western Asia, but its collection has been
mistaken for C. cerina. We have confirmed it
from Austria, Bulgaria, the Czech Republic,
Germany, Greece, Hungary, Italy, Russia,
Slovakia, Slovenia, Spain, Sweden, Turkey,
Ukraine and the United Kingdom.
Remarks. Caloplaca monacensis is characterized by its granular thallus and usually
pruinose apothecia. The North American
C. ulmorum, a closely related taxon, is very
similar, but it usually has a more developed
areolate thallus with granules at the thallus
margins.
Nomenclatural note. The name C. monacensis was never widely used; it was adopted by
Lettau (1912), Migula (1929), Zahlbruckner
(1931) and Poelt (1969) without any new
records. The reason is that the apothecia in
the exsiccate (syntype) are mostly brownish
and only rarely with the normal yelloworange colour (see Fig. 2D). Therefore it
was originally described in the genus Pyrenodesmia. In all other aspects the specimens in
the exsiccate are morphologically completely
consistent with recent collections of the
species.
Selected specimens examined. Austria: Styria:
Schladminger Tauern, Schladming, Kocourková 900419
(PRM). Lower Austria: Mitterbach am Erlaufsee,
Neuhaus, Poelt IN10-84 (GZU). Carinthia: Koralpe,
Oberauerling, Maurer 1131 (GZU); Görtschitztal,
Lölling Graben, Hafellner JH14855 (GZU). Burgenland:
Mittelburgenland, Bernstein, Maurer 356 (GZU).
Salzburg, Kondratyuk 2936 (KHER).—Bulgaria:
Kardzhali: Eastern Rodopi Mts, Ptichar, Vondrák
JV2026 (CBFS). Burgas: Strandzha Mts, Gramatikovo,
Vondrák JV3236 (CBFS).—Czech Republic: South
Moravian Reg.: Pavlovské vrchy hills, Soutěska, Vondrák
164 (hb. Šoun). Vysočina Reg.: Nové Město na Moravě,
Kut’ák 697020 (PRM). Central Bohemian Reg.: BR
Křivoklátsko, Týřov, Palice 9151 (hb. Palice).—
Greece: Crete: Orosira Dikti Mts, Ano Viannos,
Vondrák
JV3768
(CBFS).—Hungary:
BorsodAbaúj-Zemplén: Bükk Mts, Látó-kövek, Vondrák &
Khodosovtsev JV6393 (CBFS).—Italy: Trentino-Alto
Adige: Val Venosta, Glorenza, Hafellner 61270
127
(GZU).—Russia: Orenburg: Buzuluk distr., Nat. park
“Buzulukskiy bor”, Zapovednoye, Vondrák JV7336
(CBFS).—Slovakia: Banská Bystrica Reg.: Muránska
planina Mts, Mt Šiance, Bayerová, Halda & Palice 5366,
8875 (hb. Palice).—Slovenia: Cerknica: Rakek,
Prügger, Suppan, Mayrhofer & Batič SN059.8/1
(GZU).—Spain: Extremadura: SW of Cáceres, van den
Boom & van den Boom 38821 (hb. v. d. Boom).—
Sweden: Uppland: Norrsunda par., Rosenberg, Nordin
2081 (UPS).—Turkey: Eastern Anatolia Reg.: Lake
Van, Yassıca, Vondrák JV6514 (CBFS).—Ukraine:
Crimea: Czernigiv, Khodosovtsev 3042 (KHER).—Great
Britain: England: Colchester, Crombie 380 (BM).
Exsiccata examined. Austria: Lower Austria: Rosenau,
Mt Sonntagberg, Krypt. Exs. no. 252 [as C. cerina var.
ehrharti] (PRM, BM).—Bulgaria: Haskovo: Rhodope
Mts, Rabovo, Vondrák Sel. Exs. Caloplaca. no. 11 [as
C. virescens] (CBFS).—Czech Republic: Central
Bohemian Reg.: Benešov u Prahy, Hodětice, Vězda: Lich.
Sel. Exs. no. 2469 [as C. cerina var. cyanolepra] (BM).—
Germany: Baden-Württemberg: Constance, Rabenhorst:
Lich. Eur. no. 697 [as Placodium cerinum var. ehrharti]
(BM).—Italy: Veneto: Vicenza, Trevisan: Lichenotheca
veneta no. 183 [as C. cerina var. ehrhartii] (BM).—
Slovakia: Žilina Reg.: Nı́zké Tatry Mts, Lich. Slovak.
Exs. no. 227 [as C. cerina] (BM).
Caloplaca pinicola H. Magn.
Bot. Not. 1953(2): 188 (1953); type: USA, Arizona,
[Coconino Co.] Grand Canyon National Park,
Coconino Plateau, August 1926, E. & G. Du Rietz
(S!—holotype; UPS!—isotype selected by Wetmore
2004 as superfluous lectotype).
Taxonomic note. Detailed descriptions are
given in Wetmore (2004, 2007a, b). According to specimens of C. pinicola seen from
ASU, GZU and MIN, the species seems
fairly variable in thallus morphology, especially in the number and characteristics of
the soralia. The uniting character is a relatively thin spore septum (3·0–4·0 m and
never up to 5·5 m). Unfortunately, the type
specimen has a poorly developed thallus.
The species needs further study.
There is one closely related species, also
from North America, here provisionally
called C. aff. pinicola (Fig. 1), but it differs in
having larger ascospores with thicker septa.
Nomenclatural note. The holotype of C.
pinicola was identified by Magnusson himself
in S; on the label is written in Magnusson’s
handwriting “Caloplaca pinicola H. Magn. n.
sp.” and “Typus!”. Wetmore’s lectotypification (Wetmore 2004) is superfluous.
128
THE LICHENOLOGIST
Vol. 43
F. 3. Caloplaca sterilis. A, specimen with white pruinose areoles from steppe (CBFS JV6962); B, fertile specimen
without pruina from coastal sand dunes (CBFS JV7421). Scales: A & B = 1mm.
Selected specimens examined. USA: Arizona: Coconino
Co., Grand Canyon Nat. Park, Nash 10513 (GZU);
Gila Co., Flowing Springs, Nash 39318 (MIN); Pima
Co., Saguaro Nat. Mon., Wetmore 54893 (MIN).
Colorado: San Juan Co., Mesa Verde Nat. Park, Nash
18141 (ASU); Rio Grande Co., South Fork, Wetmore
17214 (MIN). New Mexico: Catron Co., San Francisco
Mts, Nash 22562 (ASU); San Juan Co., Marsh &
Rankert 419, 448 (ASU); Union Co., Capulin Mt. Nat.
Mon., Wetmore 17433 (MIN). North Dakota: McKenzie
Co., T. Roosevelt Nat. Park, Warford City, Wetmore
80071 (MIN); Billings Co., T. Roosevelt Nat. Park,
Medora, Wetmore 45047 (MIN), 80302 (GZU).
Caloplaca sterilis Šoun, Khodosovtsev,
Vondrák sp. nov.
Thallus areolatus albido-pruinosum, areolae soraliis
marginalibus; soredia viridi-grisea, pigmentis anthraquinoneis et Sedifolia-grisea in thallo et soraliis nullis;
apothecia lecanorina (Caloplaca cerinae-typus).
Typus: Ukraine, AR Crimea, Chornomorskiy district, village Olenevka, cape Tarchankut, coast of Black
Sea, on twigs of Artemisia lerchiana, 45°20#61·7$N,
32°30#47·6$E, alt. 15 m, 4 May 2010, A. Khodosovtsev
(KHER—holotypus; KHER, CBFS —isotypi).
(Fig. 3A & B)
Thallus of scattered areoles/squamules to
thin, inconspicuous, grey-green, often white
pruinose, sorediate; areoles or squamules
(50–) 157 ± 66 (–300) m wide (n = 22),
slightly convex, (50–) 85 ± 22 (–130) m
thick (n = 20); cortex alveolate (sensu Vondrák
et al. 2009a) 8–12 m thick, loosely paraplectenchymatous with crystals dissolving in
N; algal layer thick, reaching lower thallus
surface; medulla not always conspicuous,
formed by loose prosoplectenchymatous tissue; photobiont trebouxioid; soralia developed
on the areole margins or rarely punctiform on
thallus surface, greenish or greenish grey,
(80–) 145 ± 44 (–250) m in size (n = 20);
sterile thalli sometimes forming continuous
greenish sorediate crusts; soredia (15–)
24 ± 6 (–35) m diam. (n = 30), sometimes
forming consoredia (50–) 70 ± 15 (–100)
(n = 20), K−; prothallus not seen.
Apothecia lecanorine, very rare, (0·2–)
0·3 ± 0·1 (–0·6) mm diam. (n = 29), constricted at the base; disc flat to slightly convex, yellow-orange to orange, sometimes
pruinose, especially when young, K+ purple;
true exciple indistinct or very rarely visible in
section as 2–3 rows of radiate hyphae; thalline
exciple whitish pruinose, sometimes sorediate, raised above disc, (50–) 87 ± 42 (–200)
m thick (n = 20); cortex of thalline exciple
well-developed, (12–) 23 ± 9 (–40) m thick
(n = 21), paraplectenchymatous; cells (2·5–)
3·1 ± 0·5 (–4·3) m thick (n = 20); epihymenium orange, granular inspersed; hymenium 65–75 m tall, hyaline; hypothecium
50–75 m thick, hyaline; paraphyses simple to
branched, 2·0–2·5 m wide with upper cells
widened to (2·8–) 3·4 ± 0·4 (4·3) m (n =
20); asci 8-spored, (33–) 40·5 ± 3·2 (–46) ×
(10–) 13 ± 2·4 (–20) m (n = 20); ascospores
polarilocular, wide ellipsoid, rarely subglobose, (7·5–) 10·4 ± 2·0 (–16.0) × (7·5–)
7·9 ± 1·0 (–11·3) m (n = 32), septum
(2·3–) 4·7 ± 1·1 (–7·0) m wide (n = 32),
2011
The Caloplaca cerina group in Europe—Šoun et al.
ratio of septum width/spore length (0·25–)
0·5 ± 0·1 (–0·5).
Pycnidia immersed, inconspicuous; conidia
bacilliform, c. 3 × 1 m.
Etymology. The name reflects the often
sterile state of this species.
Habitat and distribution. Caloplaca sterilis
grows on basal parts of woody plant stems
(Artemisia, Halocnemum, Thymus), plant
debris and dead wood in steppe and sand
dune habitats in the Black Sea region. It is an
inconspicuous but common species in steppe
landscapes and in coastal habitats on sand
dunes in SE Europe. Associated species are
Caloplaca raesaenenii Bredkina (= C. thuringiaca Søchting & Stordeur), C. phlogina (Ach.)
Flagey, C. skii ad int., Lecania cyrtella (Ach.)
Th. Fr., Lecanora hagenii (Ach.) Ach., Physcia adscendens (Fr.) H. Olivier, Rinodina pityrea Ropin & H. Mayrhofer and Xanthoria
parietina (L.) Beltr. Hitherto known from
Bulgaria, Romania, SW Russia and Ukraine.
Two old specimens from old palings from
Cherry Hinton near Cambridge (UK)
deposited in BM (Larbalestier Lichen-Herb.
no. 167, sub Lecanora cerina var. dispersa
Larbal., 9/1879; Ibid., sub Lecanora cerina
var. cyanolepra), seem to be identical to this
species. However, molecular data is needed
to confirm the identity of British material.
Remarks. The species is characterized
by commonly white-pruinose minute
squamules/areoles with contrasting pale
greyish green to greenish soredia in punctiform, marginal or diffuse soralia. However,
our specimens from coastal sand dunes lack
pruina and their thalli do not contrast with
soralia. The colour of soredia is unique
within the C. cerina group as they do not
contain Sedifolia-grey. The species is easily
overlooked and hardly identifiable when
completely sorediate and sterile.
Specimens examined. Bulgaria: Burgas Prov.: Burgas,
Sozopol, sand dunes near seashore c. 5·5 km S of town,
on stems of Artemisia, 2007, Vondrák JV6954, JV7421,
JV7624 (CBFS).—Romania: Tulcea Co.: Babadag,
limestone outcrops 250 m SE of Enisala castle ruin, on
old dead wood in steppe, 2007, Vondrák JV6955
129
(CBFS); coast c. 5·5 km E of Sinoie, on stems of steppe
shrubs, 2007, Vondrák in JV6616 (CBFS); Razim Lake,
Popina Island, on dead twig of shrub, 2007, Vondrák
JV6963 (CBFS).—Russia: Krasnodar krai: Taman
Peninsula, steppe near road E of Primorskiy, on shrub
twig, 2007, Vondrák & Šoun JV7386 (CBFS).—
Ukraine: Kherson Reg.: Belozerskiy district, vill.
Sofievka, Sofievska balka, on Artemisia twigs, 2009,
Gavrilenko & Khodosovtsev JV6962 (CBFS), Vondrák
JV7100 (CBFS, KHER); vill. Kirove, steppe slope,
11 m alt., N46°53#92.8$ E32°50#54.9$, 24 iv 2010,
Khodosovtsev (KHER); Beryslavskiy district, village
Novokairy, Kamenska balka valley, steppe slope with
limestone outcrops, on twigs of Thymus dimorphus, alt.
20 m, 1 iv 2009, Khodosovtsev (KHER); Golopristansky
district, Black Sea reserve, Tendrivs’ka kosa island, sand
littoral dunes, on plant debris of Artemisia, 30 iv 2009,
Khodosovtsev (KHER); Gornostaivskiy district, vill.
Kairy, Kairska balka, steppe slopes, on plant debris, 21
iii 2010, Khodosovtsev (KHER); Verhnerogacheskiy district, vill. Ushkalka, cliff to Kahovskoye artificial sea,
on plant debris, 27 iii 2010, Khodosovtsev (KHER);
Novotroitskiy district, 4 km S of vill. Drushelyubovka,
cape Kutara, on coast of Sivash lake, N46°08#64.0$
E34°02#76.5$, 25 iv 2010, Khodosovtsev (KHER). AR
Crimea: Leninskiy district, Arabatskaya strelka peninsula, Valok, sand littoral dunes, on Thymus, 10 viii 2009,
Khodosovtsev (KHER); Krasnoperekopsk, Nadezdino,
at small shallow gulf S of village, on stems of Halocnemum strobilaceum, 8 vi 2006, Vondrák in JV5164
(CBFS); Pervomaiskiy district, upland Sary-Bash,
5 km SW from vill. Vypasne, near road H05, 71 m
alt., N45°35#37.9$ E33°51#49.8$, 28 iv 2010,
Khodosovtsev (KHER).
Caloplaca stillicidiorum s. lat.
Including Caloplaca stillicidiorum (Vahl) Lynge, C. cerina
var. chloroleuca (Sm.) Th. Fr., C. cerina var. muscorum
(A. Massal.) Jatta, C. cerina (Hedw.) Th. Fr. p.p., & C.
muscorum (A. Massal.) M. Choisy & Werner.
This taxon comprises Caloplaca cerina-like
lichens growing on or near the ground on
various substrata such as bryophytes, plant
debris, wood, bark of exposed roots and
chamaephytes, rarely rocks. Such specimens
form a monophyletic group in the cladogram. The molecular topology shows basically four monophyletic groups. These
groups may be only partly interpreted on the
basis of their morphology and ecology. Six
provisional ‘ecotypes’, more or less reflecting
the phylogeny (Fig. 1), are distinguished here
but they are not always separable by morphology. Characteristics of the ecotypes based on
sequenced material are briefly presented
here.
130
THE LICHENOLOGIST
1. On bryophytes and plant debris, in uplands and mountains on limestone in
Austria, Bulgaria, Czech Republic,
France, Greece, Hungary, Spain and
Ukraine; apothecia to 1·1 mm diam.;
apothecial margin strongly white pruinose,
disc sordid yellow-orange, sordid yellowgreen to sordid grey-brown, pruinose or
rarely not; thallus consisting of small white
pruinose areoles; ascospores (12–) 16·0 ±
1·8 (–19) × (6–) 8·7 ± 1·0 (–11) m, septum (5–) 7·1 ± 1·1 (–9) m wide (n = 59).
The type of Lichen chloroleucus Sm.
(BM!—holotype) and the epitype of Lichen
stillicidiorum
Vahl
(FH!—isoepitype)
selected by Wetmore (2007a) probably
belong to this ecotype. However, the holotype illustration of Lichen stillicidiorum in
Vahl (1792: 6, tab. 1063, f. 2) may belong
elsewhere.
2. On plant debris (e.g. Saxifraga) and moss,
in alpine calcareous habitats in Austria,
Russia and Sweden; apothecia to 1·2 mm
diam.; apothecial margin dark grey, white
pruinose; disc (sordid) yellow to orange,
pruinose or not; thallus invisible; ascospores (13–) 15·2 ± 1·4 (–19) × (6–)
7·7 ± 0·8 (–10) m, septum (5–) 6·3 ± 0·9
(–8) m wide (n = 43).
3. On plant debris and moss, in artic areas in
Alaska, Greenland, Norway and Svalbard;
apothecia to 1·3 mm diam.; apothecial
margin dark grey, white pruinose or not,
disc yellow-orange to orange, not pruinose;
thallus invisible; ascospores (12–) 14·7 ±
1·2 (–17) × (7–) 8·1 ± 0·7 (–9) m, septum
(5–) 5·7 ± 0·7 (–7) m wide (n = 39).
4. On plant debris, exposed wood of roots,
and rock in calcareous xerothermic habitats
in Canada, Czech Republic, Kazakhstan,
Ukraine and USA and on wooden fences in
mountains in Austria; apothecia to 1·5 mm
diam.; apothecial margin pale to dark grey,
white pruinose or not, disc yellow-orange
to orange, pruinose or not; thallus invisible
or formed by white pruinose tiny areoles;
ascospores (12–) 14·4 ± 0·9 (–16) × (5–)
7·7 ± 0·6 (–9) m, septum (4–) 4·9 ± 0·5
(–6) m wide (n = 58).
5. On acrocarpous mosses (esp. Grimmia),
on basic siliceous and conglomerate rocks
Vol. 43
in Finland, Romania, Spain, Turkey,
Ukraine, USA and Uzbekistan; apothecia
to 1·6 mm diam.; apothecial margin pale
to dark grey, white pruinose; disc yelloworange to dull orange, pruinose or not;
thallus thin to thick crustose or formed by
well-developed small lobules; ascospores
(11–) 14·8 ± 1·6 (–19) × (6–) 8·7 ± 1·3
(–11) m, septum (4–) 5·2 ± 0·6 (–7) m
wide (n = 69).
6. On bark of chamaephytes (especially
Thymus) and exposed roots, in steppelike habitats on limestone in the Czech
Republic, Hungary and Ukraine; apothecia to 0·9 mm diam.; apothecial margin
pale to medium grey, white pruinose or
not; disc yellow to yellow-orange, pruinose
or not; thallus invisible or formed by small
areoles; ascospores (10–) 12·8 ± 1·2
(–16) × (6–) 7·1 ± 0·7 (–8) m, septum
(4–) 4·8 ± 0·4 (–5) m wide (n = 39).
Burgaz et al. (1999) dealt with samples of
C. cerina on Dryas octopetala from the
Pyrenees with extremely wide septa, 7–9 m,
more than half of the length of the ascospore.
Such wide septa have not been observed in
the material studied here and in future attention should be given to those samples.
Caloplaca subalpina Vondrák, Šoun &
Palice
Lichenologist 40: 378 (2008); type: Ukraine, Eastern
Carpathians, Svidovets Mts, glacial cirque in NE
slope below Mt Bliznitsa, alt. c. 1500 m, 48° 14# 21$
N, 24° 14# E, on lime-rich schist outcrop, beneath
overhang, in subalpine belt, 29 June 2007, Vondrák
CBFS JV6072 (CBFS!—holotype; GZU, L—isotypes).
This saxicolous species grows on well-lit
rocks beneath overhangs and is known from
several localities throughout the mountains
in Europe. Apothecia are rarely developed on
the white-pruinose areolate thallus which
merges into squamules at margins. The soralia are dark grey, arising from margins
of areoles. For a detailed description see
Vondrák et al. (2008).
Caloplaca thracopontica Vondrák & Šoun
Lichenologist 40: 381 (2008); type: Turkey, Black Sea
coast, Sinop, coastal rocks on NE coast of peninsula, alt.
2011
The Caloplaca cerina group in Europe—Šoun et al.
c. 100 m, 42° 01# 57·81$ N, 35° 11# 34·42$ E, on
coastal volcanic rock, 21 April 2007, Vondrák CBFS
JV5419 (CBFS!—holotype; GZU, hb. M. Seaward—
isotypes).
This is a maritime saxicolous species that
grows on hard siliceous outcrops in the Black
Sea region, where it is locally common.
It is characterized by the presence of
lobules and pustules on the thallus. On the
Mediterranean coasts there is a similar, but
possibly different species, hitherto not well
known (see Results). For a detailed description see Vondrák et al. (2008).
Caloplaca turkuensis (Vain.) Zahlbr.
Cat. Lich. Univ. 8: 589 (1931) [as ‘turkensis’].—
Placodium turkuense Vain., Ann. Soc. zool.-bot. Fenn.
Vanamo 9: 320 (1929); type: Ab. Turku, Vartiovuori, 2
Oct. 1927, Vainio 35118 (TUR-V 35374!—holotype).
Caloplaca jemtlandica H. Magn., Bot. Notiser 1945:
304 (1945); type: Sweden, Jämtland, Undersåker, 30 vi
1912, Malme (S!—holotype, UPS!—isotype).
Caloplaca jemtlandica var. cerinosora E. S. Hansen,
Poelt & Søchting, Meddelelser om Grønland, Bioscience
25: 33 (1987); type: Grönland, Disko, Lyngmark, untere Hänge des Lyngmarksfjeld N Godhavn, an Salix, 27
vii 1983, Poelt & Ullrich (GZU!—holotype).
Caloplaca cerina f. coronulata Th. Fr., Lichenogr.
Scandinavica I: 175 (1871).—Lecanora cerina f. coronulata Nyl., Notiser ur Sällsk. pro Fauna et Flora Fennica
Förhandlingar 8: 128 (1866); type: Lapponia orientalis.
Knäsä [Russia, Murmansk Oblast, Knyazhaya Guba],
1861, Fellman (H!—lectotype designated here).
(Figs 2E & F)
Thallus with very small areoles, sorediate,
up to several cm wide, non-pruinose, up to
0·13 (–0·3) mm thick; areoles sordid white to
pale (sordid) grey, initially convex but later
flat, up to 0·2 mm wide, usually very soon
entirely sorediate. Soralia pale to dark grey or
greenish grey, with bluish tinge, whitish or
greenish inside (seen when external soredia
are detached), arising from upper surface
or margins of areoles, scattered to confluent, slightly convex; soredia (15–) 22 ± 3
(–30) m diam. (n = 95); prothallus sometimes
present, very thin, grey to black; cortex up to
10 (–20) m thick, paraplectenchymatous.
Apothecia lecanorine, abundant to absent,
up to 0·8 mm diam., sessile; disc flat to
slightly convex, yellow-orange, rarely pruinose; true exciple indistinct, up to 15 m
131
thick, prosoplectenchymatous; thalline exciple
grey to white, rarely pruinose, raised above
the disc when young, somewhat reduced in
old apothecia, 50–85 m thick, sometimes
becoming sorediate; epihymenium orange,
with inspersed granules; hymenium 55–
100 m thick, hyaline; hypothecium 55–
110 m thick, hyaline, sometimes oilinspersed; paraphyses simple to branched,
2 m broad with upper cells wider, 3–5 m
diam.; asci 8-spored, 38–54 × 10–17 m
(n = 19); ascospores polarilocular, ellipsoid,
(10·0–) 12·1 ± 1·1 (–15·0) × (6·0–) 6·9 ±
0·8 (–9·0) m, septum (3·0–) 4·3 ± 0·7
(–6·0) m wide (n = 58), ratio of septum
width/spore length (0·2–) 0·4 ± 0·05 (–0·5).
Pycnidia sometimes present, immersed,
distinct by elevated darker ostiole; conidia
bacilliform, 3–4 × 1 m.
Habitat and distribution. This species grows
on bark of broad-leaved trees (Acer, Alnus,
Betula, Fraxinus, Malus, Populus, Sorbus,
Ulmus) and rarely of conifers (Picea). Also
found on wood of spruce stumps and
wooden fences. It occurs from lowlands to
mountains in deciduous and mixed forests
or on wayside trees. The specimen from
Greenland (C. jemtlandica var. cerinosora)
grew on a twig of Salix glauca. Probably, this
species is widespread throughout Europe
and North America, but it has not previously been recognized. Hitherto recorded
from Austria, Czech Republic, Finland,
Greenland, Norway, Russia, Slovakia,
Sweden, United Kingdom and the USA.
Remarks. This species is characterized by
the often sorediate apothecial margins,
slightly convex, punctiform to confluent
blue-grey soralia, minute or often absent
areoles. However, specimens without sorediate apothecial margins and with persistent
corticated areoles with delimited soralia are
common and can be confused with small
thalli of C. chlorina, but the latter usually has
larger soredia. Sterile specimens can be confused with Rinodina colobina (Ach.) Th. Fr.
or Rinodina pityrea Ropin & H. Mayrhofer.
During the phylogenetic analyses a taxon
similar to C. turkuensis was discovered, but
132
THE LICHENOLOGIST
several ITS sequences showed that it
belonged outside of the C. cerina group (not
included in the cladogram in Fig. 1). These
specimens from Sweden are morphologically
distinguished from C. turkuensis by soralia
and apothecia with an endophloeic origin,
not developing from areoles, the true exciple
is usually more apparent and by the thalline
margin being not so stout. This is probably
an unknown species which needs further
study. The types of C. turkuensis and the
here synonymized C. jemtlandica incl. var.
cerinosora and C. cerina f. coronulata are
mostly old and were not included in the
molecular analysis. However, morphologically they seem to correspond with our
material molecularly confirmed to belong to
the C. cerina group.
Nomenclatural note. Specimens of this
species have been commonly misidentified as
various species, i.e., C. virescens, C. chlorina
or C. cerina. However, the names Caloplaca
jemtlandica (including var. cerinosora) and
Caloplaca turkuensis have been applied to different forms of the same species, but the
latter name is the oldest and has therefore
priority over the younger ones.
Caloplaca turkuensis has probably been
used only for the type collection, which has a
fairly well-developed, thick, sorediate thallus
and abundant apothecia, with completely
sorediate margins; it is congruent in phenotype with specimens used for obtaining the
molecular data.
Caloplaca jemtlandica is also a little known
taxon. Surprisingly, the type specimens are
sorediate, contrary to the description in the
original paper, although both thallus and
apothecial margin are not so sorediate as in
the type of C. turkuensis. The specimen
labelled as isotype in S is not conspecific with
holotype in S and isotype in UPS, but at the
moment it is not possible to determine where
it belongs.
Caloplaca jemtlandica var. cerinosora is
another poorly known taxon, described and
hitherto known only from one twig of
Salix glauca from Greenland. The type specimen is a poorly developed specimen of C.
turkuensis.
Vol. 43
Caloplaca cerina f. coronulata is a form with
distinctly sorediate thallus and apothecial
margin clearly conspecific with C. turkuensis.
Selected specimens examined. Austria: Vorarlberg:
Verwall-Gruppe, Nenzigast-Tal, Poelt IN85-86 (GZU).
Styria: Schladming, Aich, Vondrák JV7251 (CBFS).—
Czech Republic: Central Bohemian Reg.: Přı́bram,
Lešetice, Malíček et al. 1258 (hb. Malı́ček). Plzeň Reg.:
Šumava Mts, Modrava, Mt Medvěd, Liška JV2921
(CBFS). South Bohemian Reg.: Šumava Mts, České
Žleby, Palice, Dětinský, Horáková & Liška (hb. Palice).
Moravian-Silesian Reg.: Beskydy Mts, Staré Hamry,
Salajka, Malíček & Palice 1908 (hb. Malı́ček).—
Finland: Uusimaa: Karjalohja, Maila, Laurila (H).
Tavastia Proper: Tammela, Mustiala, Räsänen (H).—
Norway: Oslo, Tønsberg 9926 (BM). Buskerud: Sigdal,
Prestfoss, Tønsberg 13312 (BM). Oppland: Vang, Lykkja,
Søchting (BM).—Russia: Orenburg: Buzuluk distr., Nat.
park “Buzulukskiy bor”, Zapovednoye, Vondrák JV7429
(CBFS).—Slovakia: Prešov Reg.: E Carpathians, N.P.
Nı́zké Poloniny, Mt Pl’aša, Palice & Šárová 9302
(PRA).—Sweden: Torne Lappmark: Jukkasjärvi par.,
Torneträsk, Arup L04229 (LD). Gotland: Atlingbo par.,
Lillo Atlings, Sundin & Thor 57 (UPS).—United
Kingdom: Scotland: Elgin, Ballindalloch Castle, James
(BM).—USA: Washington: Pend Oreille Co., Selkirk
Mts, Lower Granite Falls, Spribille 15393 (hb. Spribille).
Caloplaca ulmorum (Fink) Fink
Lichen Fl. U. S. 358 (1935).—Placodium cerinum var.
ulmorum Fink, Contr. U. S. Nat. Herb. 14: 215 (1910);
type: USA, Kansas, Rooks County, on Juniperus virginiana, 7 xi 1893, E. Bartholomew (MICH 00062622!—
lectotype designated by Wetmore 2007a; MICH
00067693!—isolectotype).
Thallus areolate to predominantly granular, up to 0·16 (–0·3) mm thick; areoles flat to
slightly convex, scattered to continuous,
non-pruinose, beige, beige-grey to pale
greenish grey, up to 0·8 mm wide; granules
(50–) 107 ± 32 (–200) m diam. (n = 90);
cortex up to 25 (–40) m thick, paraplectenchymatous, sometimes made very minutely
tomentose by inconspicuous hairs 4–10 ×
2–3 m formed by projecting hyphae (visible
in section), particulary on granules; prothallus
not seen.
Apothecia lecanorine, usually abundant, up
to 1·3 mm diam., sessile, distinct, usually
pruinose; disc flat to slightly convex, yellow to
yellow-orange, usually pruinose; true exciple
indistinct, very thin, up to 10 m thick, prosoplectenchymatous; thalline exciple of thallus
colour, usually white pruinose, raised above
2011
The Caloplaca cerina group in Europe—Šoun et al.
disc when young, somewhat reduced in old
apothecia, 75–120 m thick; epihymenium
orange, with inspersed granules; hymenium
80–100 m thick, hyaline; hypothecium 50–
140 m thick, hyaline; paraphyses simple to
branched, 2 m broad with upper cells wider,
3–6 m thick; asci 8-spored, 42–70 × 10–18
m (n = 45); ascospores polarilocular, ellipsoid, (11·0–) 13·6 ± 1·4 (–18·0) × (6·0–)
6·9 ± 0·8 (–9·0) m; septum (4·0–) 5·6 ±
0·9 (–8·0) m wide (n = 90); ratio of septum
width/spore length (0·3–) 0·4 ± 0·05 (–0·5).
Pycnidia rare to abundant, immersed,
distinct by elevated darker ostiole; conidia
bacilliform, 3–4 × 1 m.
Habitat and distribution. Caloplaca ulmorum
is an exclusively North American species
usually growing on bark of broad-leaved
trees, mainly Ulmus but also on, for example,
Acer and Quercus. Nevertheless, the type collection is from bark of Juniperus virginiana.
The complete distribution is not known.
133
Taxa with uncertain affinities
Caloplaca virescens (Sm.) Coppins seems
to be morphologically and ecologically close
to C. monacensis, but differs in its welldeveloped, areolate, rarely fertile thallus with
a thick layer of small granules (45–70 m
diam.). Unfortunately, no fresh material was
available for sequencing. Four very old specimens, including the part of the holotype (see
Laundon 1992), and only one more recent
specimen (1973) have been found in BM.
This species needs further study.
The little used name C. albolutea (Nyl.)
Zahlbr. (type: Supra muscos pulvinatos in
Lapponia orientali eam in insulis Maris Albi,
legit G. Selin, H-NYL!—lectotype designated here) and perhaps C. mydalaea (Körb.)
Zahlbr. (type: UPS—“fragment. specimin.
original.”!) probably belong to C. stillicidiorum s. lat. but both type specimens are too
exiguous.
Discussion
Remarks. The species is characterized by a
usually well-developed areolate thallus with
marginal granules. Poorly developed thalli
consisting of only granules are morphologically hard to separate from C. monacensis. A
good illustration may be found in Wetmore
(2007a: 805).
Nomenclatural note. Wetmore (2007a)
synonymized this name with C. cerina but we
have shown that it represents a separate
species.
Selected specimens examined. USA: Minnesota:
Hennepin Co., Minneapolis, Wayzata Big Woods,
Wetmore 90256 (GZU), ibid., Wolsfeld Woods, Trana
9973 (GZU); Olmsted Co., Oronoco, Wetmore 24185
(GZU). Missouri: Newton Co., Joplin, George
Washington Carver Nat. Mon., Wetmore 69226 (LD).
South Dakota: McCook Co., Lake Vermillion St. Rec.
Area, Wetmore 93154 (MIN). Wisconsin: Bayfield Co.,
Rainbow Lake Wilderness, Bellevue Lake, Wetmore
70829 (GZU).
Exsiccata examined. Canada: Ontario: Ontario Co.,
Beaverton, Lich. Can. Exs. no. 157 (GZU).—USA:
South Dakota: Grant Co., Marvin, Blue Cloud
Benedictine Abbey, Lich. E. N. Am. Exs. no. 265
[as Caloplaca cerina] (GZU). Wisconsin, Iowa Co.,
Barneveld, Lich. Wiscon. Exs. no. 107 (GZU).
Our results show again, what has already
been proved many times; the phenotypebased traditional taxonomy has been influenced by choosing incorrect diagnostic
characters or by the incorrect interpretation
of the characters. For example, in the British
lichen flora, Fletcher & Laundon (2009) list
three species of the Caloplaca cerina group:
a sorediate “C. virescens”, an isidiate “C.
chlorina” and “C. cerina” without vegetative
diaspores. This is a very practical simplification, but the situation is much more complex. As we have shown, sorediate and
isidiate species and species without vegetative diaspores occur repeatedly within the
phylogenetic tree, and very similar species
are often not closely related. Based on our
data, we cannot say which thallus characters
are plesiomorphic and which are derived,
but convergent evolution is evident here in
phenotypes even with or without vegetative
diaspores. Similar patterns have been
observed in the phylogeny of the C. citrina
group (Arup 2006, Vondrák et al. 2009a) and
the C. ferruginea group (Arup & Åkelius
2009).
134
THE LICHENOLOGIST
Our molecular analysis corroborates the
sometimes underestimated role of the precise
type of vegetative diaspores as a crucial
phenotypic character in this complex. For
instance, presence of soredia (or blastidia) vs.
isidia clearly separate the formerly often
unrecognized C. chlorina and C. isidiigera.
Undoubtedly, other important phenotype
characters are substratum specificity and
geographical distribution. This is shown in
a provisional taxonomy of C. cerina s. lat.
and C. stillicidiorum s. lat., where vegetative
diaspores are absent, and only a little data on
ecology and distribution are available. Both
groups are formed of several phylogenetic
lineages, polyphyletic in C. cerina s. lat. and
monophyletic in C. stillicidiorum s. lat., which
may be called cryptic species or semi-cryptic
species, if they have at least some phenotypic
tendencies (Vondrák et al. 2009a).
Grube & Kroken (2000) recommended
that a phylogenetic species should be congruent with at least one phenotypic character
when a single locus such as ITS is used.
Crespo & Pérez-Ortega (2009) suggested for
recognition of cryptic species that molecular
analysis should be based on more than one
independent genetic region, in order to detect recombination events among lineages.
We are so far not able to fulfill these conditions in the two species complexes. Several
loci, more extensive sampling to avoid
present sampling bias and/or more advanced
phenotype appraisal should be used to separate some putative cryptic species within these
tentative aggregate names.
We are grateful to Z. Palice, P. van den Boom, T.
Spribille, J. P. Halda, J. Malı́ček and to the following
herbaria and their curators: B (H. Sipman), C (E. S.
Hansen), FH (M. Schmull), GZU (W. Obermayer),
ASU (T. H. Nash III), BM (H. Thüs), H (L. Myllys), M
(A. Beck), MIN (I. Schmitt), PRM (J. Kocourková),
TUR (S. Huhtinen), UPS (A. Nordin) for loaning us
their herbarium material. J. Košnar kindly collected
some fresh material during his field trip to Norway and
Svalbard. O. Redchenko kindly made some sequences.
M. Herbstová (University of South Bohemia), L.
Knudsen and A. Løth (University of Copenhagen)
kindly helped JŠ in the molecular laboratories. Our work
was financially supported by the Grant Agency of the
Academy of Sciences of the Czech Republic (KJB
601410701) and Ministry of Education of the Czech
Republic grant no. MSM 6007665801. The work in
Vol. 43
Copenhagen was supported by a grant from the European Commission’s (FP 6) Integrated Infrastructure
Initiative programme SYNTHESYS (DK-TAF 2664
and 4724).
R
Arup, U. (2006) A new taxonomy of the Caloplaca citrina
group in the Nordic countries, except Iceland.
Lichenologist 38: 1–20.
Arup, U. & Åkelius, E. (2009) A taxonomic revision of
Caloplaca herbidella and C. furfuracea. Lichenologist
41: 465–480.
Burgaz, A. R., Arenal, F., González, V. &
Esteve-Raventós, F. (1999) Caloplaca cerina in the
Pyrenees mountains of Huesca Province. Botanica
Complutensis 23: 53–57.
Clauzade, G. & Roux, C. (1985) Likenoj de Okcidenta
Europo. Ilustrita Determinlibro. Bulletin de la
Société Botanique du Centre-Ouest, Nouvelle Série,
Numéro Spécial 7: 1–893.
Crespo, A. & Pérez-Ortega, S. (2009) Cryptic species
and species pairs in lichens: a discussion on the
relationship between molecular phylogenies and
morphological characters. Anales del Jardín Botánico
de Madrid 66S1: 71–81.
Ekman, S. (2001) Molecular phylogeny of the
Bacidiaceae (Lecanorales, lichenized Ascomycota).
Mycological Research 105: 783–797.
Fletcher, A. & Laundon, J. R. (2009) Caloplaca. In The
Lichens of Great Britain and Ireland. (C. W. Smith, A.
Aptroot, B. J. Coppins, A. Fletcher, O. L. Gilbert,
P. W. James & P. A. Wolseley, eds): 245–273.
London: The British Lichen Society.
Gardes, M. & Bruns, T. D. (1993) ITS primers with
enhanced specificity for basidiomycetes. Application for the identification of mycorrhizae and rust.
Molecular Ecology 2: 113–118.
Grube, M. & Kroken, S. (2000) Molecular approaches
and the concept of species and species complexes
in lichenized fungi. Mycological Research 104:
1284–1294.
Hansen, E. S., Poelt, J. & Søchting, U. (1987) Die
Flechtengattung Caloplaca in Grönland. Meddelelser
om Grønland, Bioscience 25: 1–52.
Hedwig, J. (1788) Descriptio et Adumbratio MicroscopicoAnalytica Muscorum Frondosorum 2. Mülleriano,
Lipsia.
Kärnefelt, I. (1990) Isidiate taxa in the Teloschistaceae
and their ecological and evolutionary significance.
Lichenologist 22: 307–320.
Kärnefelt, I. & Kondratyuk, S. Y. (2004) Contributions
to the lichen genus Caloplaca (Teloschistaceae) from
Australia. Bibliotheca Lichenologica 88: 255–265.
Katoh, K., Kuma, K., Toh, H. & Miyata, T. (2002)
MAFFT: a novel method for rapid multiple
sequence alignment based on fast Fourier transform. Nucleic Acids Research 30: 3059–3066.
Laundon, J. R. (1992) Lepraria in the British Isles.
Lichenologist 24: 315–350.
Lettau, G. (1912) Beiträge zur Lichenographie von
Thüringen. Hedwigia 52: 81–264.
2011
The Caloplaca cerina group in Europe—Šoun et al.
Meyer, B. & Printzen, C. (2000) Proposal for a standardized nomenclature and characterization of
insoluble lichen pigments. Lichenologist 32:
571–583.
Migula, W. (1929) Die Flechten. In Thomé-Migula,
Flora von Deutschland, Österreich und der Schweiz.
Abt. II. Kryptogamen-Flora, Band XII/1, BerlinLichterfelde: H. Bermühler.
Nylander, J. A. A. (2004) MrModeltest v2. Program
distributed by the author. Evolutionary Biology
Centre, Uppsala University.
Poelt, J. (1969) Bestimmungsschlüssel Europäischer
Flechten. Lehre: J. Cramer.
Posada, D. & Crandall, K. A. (1998) MODELTEST—
testing the model of DNA substitution. Bioinformatics 14: 817–818.
Rodriguez, F., Oliver, J. L., Marin, A. & Medina, J. R.
(1990) The general stochastic model of nucleotide
substitution. Journal of Theoretical Biology 142:
485–501.
Ronquist, F. & Huelsenbeck, J. P. (2003) MrBAYES 3:
Bayesian phylogenetic inference under mixed
models. Bioinformatics 19: 1572–1574.
Søchting, U. (1997) Two major anthraquinone chemosyndromes in Teloschistaceae. Bibliotheca Lichenologica 68: 135–144.
Vahl, M. (1792) Flora Danica, Issue 18, Table 1063,
Fig. 2.
van den Boom, P. P. G. & Rico, V. J. (2006) Caloplaca
squamuloisidiata, a new lichen from Portugal and
Spain. Lichenologist 38: 529–535.
Vondrák, J., Šoun, J., Hrouzek, P., Řı́ha, P., Kubásek,
J., Palice, Z. & Søchting, U. (2008) Caloplaca subalpina and C. thracopontica, two new saxicolous
135
species from the Caloplaca cerina group (Teloschistaceae). Lichenologist 40: 375–386.
Vondrák, J., Řı́ha, P., Arup, U. & Søchting, U. (2009a)
The taxonomy of the Caloplaca citrina group (Teloschistaceae) in the Black Sea region; with contributions to the cryptic species concept in lichenology.
Lichenologist 41: 571–604.
Vondrák, J., Šoun, J., Redchenko, O., Lőkös, L. &
Khodosovtsev, A. (2009b) Populations of two Caloplaca species with peculiar ecology observed in the
Bükk Mts, Hungary. Bryonora 44: 8–12.
Wetmore, C. M. (1996) The Caloplaca sideritis group in
North and Central America. Bryologist 99: 292–314.
Wetmore, C. M. (1997) The typification of Caloplaca
chlorina. Bryologist 100: 170.
Wetmore, C. M. (2004) The sorediate corticolous
species of Caloplaca in North and Central America.
Bryologist 107: 505–520.
Wetmore, C. M. (2007a) Notes on Caloplaca cerina
(Teloschistaceae) in North and Central America.
Bryologist 110: 798–807.
Wetmore, C. M. (2007b) Caloplaca. In Lichen Flora of
the Greater Sonoran Desert Region. Volume 3. (T. H.
Nash III, C. Gries & F. Bungartz, eds): 179–220.
Tempe: Lichens Unlimited, Arizona State
University.
White, T. J., Bruns, T. D., Lee, S. & Taylor, J. (1990)
Amplification and direct sequencing of fungal ribosomal DNA genes for phylogenies. In PCR Protocols:
a Guide to Methods and Applications (M. A. Innis, D.
H. Gelfand, J. J. Sninsky & T. J. White, eds): 315–
322 . San Diego: Academic Press.
Zahlbruckner, A. (1931) Catalogus Lichenum Universalis.
7. Leipzig: Borntraeger.
Accepted for publication 17 October 2010