New Crustose Teloschistaceae in Central Europe

The Lichenologist 45(6): 701–722 (2013) 6 British Lichen Society, 2013 doi:10.1017/S0024282913000455

New crustose Teloschistaceae in Central Europe

Jan VONDRA´ K, Ivan FROLOV, Pavel Rˇ I´HA, Pavel HROUZEK, Zdeneˇk PALICE, Olga NADYEINA, Go¨khan HALICI, Alexander KHODOSOVTSEV and Claude ROUX

Abstract: Central Europe in general is poor in Teloschistaceae lichen crusts (Caloplaca s. lat.). Diver- sity of these lichens is increased by the occurrence of some Arctic, Mediterranean and continental species, which are here close to the limits of their range. Examples include: 1) Caloplaca interfulgens, previously known from arid territories of northern Africa and western Asia, is recorded, surprisingly, from Austria, Czech Republic, Germany, Slovakia and southern Russia. In Central Europe, it is restricted to scattered xerothermic limestone outcrops. 2) Caloplaca scabrosa, previously known only from Svalbard, is recorded from the Sudetes in the Czech Republic. It is similar to, but not conspecific with, C. furfuracea. Its diagnostic characters include a blastidiate thallus and the presence of atranorin. Our results show that atranorin is absent in the majority of taxa related to C. furfuracea with only two exceptions: the sample from Eastern Carpa- thians, here called C. aff. scabrosa, and in one Sudetan sample identified as C. crenularia. 3) Caloplaca emilii, newly described below, is closely related to the Mediterranean C. areolata.We consider C. emilii a Mediterranean species rarely occurring in higher latitudes in Austria, the Czech Re- public and Germany. It is distinguished from C. areolata mainly by the presence of vegetative diaspores (blastidia); a possible role of blastidia in the distribution pattern of C. emilii is discussed below. Status of the names Caloplaca areolata, C. isidiigera and C. spalatensisis, formerly used for the new taxon, is clarified. 4) Caloplaca molariformis, newly described below, belongs to the Pyrenodesmia group (a lineage of Caloplaca without anthraquinones). It is a continental species, frequently collected on limestone or lime-rich tuffs in steppes or deserts in Turkey, Iran, western Kazakhstan and southern Russia, and is also known from eastern Ukraine and southern Slovakia. Caloplaca molariformis is characterized by its thick thallus with fungal and algal tissues arranged in high stacks. 5) Caloplaca substerilis, newly described below, is distinguished from the closely related C. ulcerosa by its endophloeodal or minutely squamulose thallus with soralia formed in bark crevices or on margins of squamules. While C. ulcerosa has a maritime distribution in Europe, C. substerilis is typically a continental species. North American continental lichens called ‘‘C. ulcerosa’’ are phylogenetically closer and more similar to C. substerilis. The positions within Teloschistaceae of the taxa considered are demonstrated by ITS phylogenies. The distributions of C. areolata, C. emilii and C. interfulgens are mapped. The new species are fully described using more than a hundred phenotype characters, and diagnostic characters are indicated separately. Key words: biodiversity, biogeography, ITS phylogeny, lichen phenotype evaluation, species recogni- tion, vegetative reproduction Accepted for publication 26 May 2013

J. Vondra´k: Institute of Botany, Academy of Sciences, Za´mek 1, Pru˚honice, CZ-25243, Czech Republic; De- Za´mek 1, Pru˚honice, CZ-25243, Czech Republic; De- partment of Botany, Faculty of Natural Sciences, partment of Botany, Faculty of Science, University of Charles University, Bena´tska´ 2, Praha, CZ-128 01, Czech South Bohemia, Branisˇovska´ 31, Cˇ eske´ Budeˇjovice, CZ- Republic. 370 05, Czech Republic. Email: [email protected] O. Nadyeina: Department of Lichenology and Bryology, I. Frolov: Faculty of Science, University of South Bohe- M.G. Kholodny Institute of Botany, National Academy mia, Branisˇovska´ 31, Cˇ eske´ Budeˇjovice, CZ-370 05, of Sciences of Ukraine, Tereschenkivska str. 2, 01601 Czech Republic; Faculty of Biology, Ural Federal Uni- Kiev, Ukraine; Biodiversity and Conservation Biology versity, ul. Mira 19, Ekaterinburg, 620002, Russia. Swiss Federal Institute for Forest, Snow and Landscape P. Rˇ ´ıha: Department of Botany, Faculty of Science, Uni- Research, WSL Zu¨rcherstr. 111, CH-8903, Birmensdorf, versity of South Bohemia, Branisˇovska´ 31, Cˇ eske´ Budeˇ- Switzerland. jovice, CZ-370 05, Czech Republic. G. Halıcı: Erciyes University, Faculty of Science, De- P. Hrouzek: Department of Autotrophic Microorgan- partment of Biology, 38039, Kayseri, Turkey. isms, Institute of Microbiology, Academy of Sciences, A. Khodosovtsev: Kherson State University, 40 Rokiv Opatovicky´ mly´n, Trˇebonˇ, CZ-379 81, Czech Republic. Zhovtnya str. 27, 73000 Kherson, Ukraine. Z. Palice: Institute of Botany, Academy of Sciences, C. Roux: Chemin des Vignes-Vieilles, FR-84120, Mira- beau, France.

Introduction older herbarium samples from BRA and STU (in Calo- placa interfulgens and C. emilii) are as complete as we Teloschistaceae, with its 1000 or more species can make them. Specimens from CBFS, PRA and GZU (Arup et al. 2013), has highest biodiversity in used for comparative studies are cited more briefly in the temperate regions (Feuerer 2011). In Cen- text. tral Europe, hot spots of Caloplaca diversity Phenotype evaluation are restricted to habitats with sun-exposed calcareous or base-rich siliceous outcrops in More than 100 phenotype characters were assessed alpine zones of the Alps and high Carpathians before preparing descriptions of the three new taxa. The list of characters and the way in which they were studied (e.g. Poelt 1953a, b, 1954, 1955, 1960, 1964; is provided in Vondra´k et al. (2013). All observations Wilk & Flakus 2006; Vondra´k et al. 2008), or were carried out on dead, stabilized material, on hand- in dry and warm rocky steppes (e.g. Poelt cut sections mounted in water, without any chemical 1975; Vondra´k et al. 2007). In other Central treatments. Measurements are accurate to 05 mm for cells and 10 mm for larger structures. All measurements European habitats, only the common epiphytic of cells include their walls, except for tissues with gluti- and epilithic species are found; the highest nized cell walls. In Caloplaca molariformis, the widths of number of these common species is found algal and fungal stacks are measured at the mid-point on lime-rich artificial substrata (e.g. Vondra´k of their vertical extent. In each sample, ten measure- & Hrouzek 2006; Svoboda et al. 2007; Von- ments were made for each measurable character. Results of the measurements are given as (min.–) x1#–x2#–x#3 dra´k et al. 2010a). (–max.), where min/max are extremes from all meas- Altogether, more than one hundred Calo- urements, x#1 is the lowest specimen arithmetic mean ob- placa species occur in Central Europe (Von- served, x2#is the arithmetic mean of all observations, x3#is dra´k & Wirth 2013), but about two thirds of the highest specimen arithmetic mean observed. In cases these are rare species, known from very few where measurements were made from one sample, only x2#is recorded. Total number of measurements (n), localities. In other words, the generally low number of samples asessed (N), and standard deviation Caloplaca species diversity in Central Europe from all measurements (SD) are given in square paren- is partly enriched by marginal occurrences thesis for each character measured [n; N; SD]. General of some ‘exotic’ taxa further distributed in morphological terminology follows Smith et al. (2009); the term ‘‘alveolate cortex’’ is adopted from Vondra´k et the Mediterranean basin, western Asia or in al. (2009a). the Arctic. Known examples are C. exsecuta (Nyl.) Dalla Torre & Sarnth., C. haematites Chemistry (Chaub.) Zwackh, C. pollinii (A. Massal.) Spot tests with KOH (K), sodium hypochlorite (C), Jatta (Vondra´k & Wirth 2013), C. raesaenenii paraphenylenediamine (P) and UV light were performed Bredkina (e.g. Søchting & Stordeur 2001), in each new species. Tissues were also tested for amy- C. tominii Savicz (Vondra´k et al. 2011), and loidity by the reaction with Lugol’s solution (I). Pig- many others. ments insoluble in acetone were evaluated following Here we report several taxa newly discov- Meyer & Printzen (2000). Extracellular crystals were examined by the reaction with concentrated H2SO4 for ered in Central Europe. Caloplaca interfulgens detection of Ca. HPLC was used for identification of and C. scabrosa were previously known only acetone-soluble compounds. The anthraquinone contents from very distant areas and their occurrence were analyzed on a LichroCART 250-4 RP18-e (5 mm) in Central Europe was not expected. Calo- column using an Agilent 1100 Series Chromatograph after Søchting (1997), but using the wavelength (240 placa emilii, C. molariformis and C. substerilis nm). Whole absorption spectra in the range 200–600 are newly described from elsewhere, but also nm were monitored. The presence of atranorin in the occur in Central Europe. samples was determined after Feige et al. (1993) on the same column and chromatographic system.

Materials and Methods DNA extraction, ampliﬁcation and sequencing The simple NaOH extraction (Werner et al. 2002) was Sampling used for DNA isolations. Primers for PCR ampliﬁcation Lichen samples were collected by the authors from were ITS1F (Gardes & Bruns 1993) and ITS4 (White various European and Asian localities between 1994 and et al. 1990). PCR cycling parameters follow Ekman 2012. We list information regarding locality, habitat, (2001). A total of 51 nuclear ITS sequences were newly collection and deposition of specimens. Citations of the generated (Table 1).

Table 1. New Caloplaca ITS sequences generated during this study

GenBank Labels of phylogenetic tree terminals Voucher Accession No.

Caloplaca aff. crenularia Canary Islands GZU (1993, Poelt & Sanchez-Pinto) KC416116 C. areolata Croatia CBFS JV7950 KC416098 C. areolata Montenegro GZU (2008, Mayrhofer) KC416097 C. areolata Spain CBFS JV6314 KC416096 C. crenularia Bulgaria CBFS JV2065 KC416112 C. crenularia Crete 1 GZU (Mayrhofer 18045) KC416113 C. crenularia Crete 2 CBFS JV4137 KC416119 C. crenularia Hungary CBFS JV6409 KC416117 C. crenularia Iran CBFS JV5608 KC416115 C. crenularia Spain CBFS JV6255 KC416114 C. crenularia Turkey CBFS JV6064 KC416118 C. emilii Bulgaria, holotype CBFS JV6600 KC416101 C. emilii Bulgaria, Rhodopes CBFS JV2223 KC416099 C. emilii Czech Republic CBFS JV9358 KC416102 C. emilii Czech Republic 2 CBFS JV9357 KC416103 C. emilii France Hb. Clauzade 23475 KC416100 C. emilii Greece CBFS JV8832 KC416104 C. ferrarii s.lat. Czech Republic 1 CBFS JV8782 KC416139 C. ferrarii s.lat. Czech Republic 2 CBFS JV9150 KC416132 C. ferrarii s.lat. Czech Republic 3 CBFS JV9043 KC416137 C. ferrarii s.lat. Czech Republic 4 CBFS JV9151 KC416140 C. furfuracea Austria PRA (Palice 12390) KC416120 C. fuscorufa Ukraine CBFS JV6204 KC416111 C. herbidella Turkey PRA (Palice 11832) KC917268 C. interfulgens Czech Republic 1 CBFS JV9399 KC416134 C. interfulgens Czech Republic 2 CBFS JV9153 KC416131 C. interfulgens Czech Republic 3 CBFS JV9156 KC416129 C. interfulgens Czech Republic 4 CBFS JV9155 KC416130 C. interfulgens Czech Republic 5 CBFS JV9144 KC416138 C. interfulgens Slovakia 1 CBFS JV9260 KC416136 C. interfulgens Slovakia 2 CBFS JV9186 KC416135 C. interfulgens southern Russia CBFS JV9396 KC416133 C. interfulgens Turkey CBFS JV8552 KC416125 C. interfulgens Turkey CBFS JV8557 KC416126 C. interfulgens Turkey CBFS JV8539 KC416127 C. lactea Greece CBFS JV8331 KC416128 C. lactea Italy CBFS JV8679 KC416124 C. molariformis Kazakhstan CBFS JV7635 KC416146 C. molariformis Slovakia, holotype CBFS JV10192 KC416142 C. molariformis Turkey CBFS JV9787 KC416144 C. molariformis Ukraine 1 KV (Luhansk, Nadyeina 132) KC416143 C. molariformis Ukraine 2 KV (Luhansk, Nadyeina 134) KC416145 C. scabrosa Czech Republic CBFS JV1908 KC416122 C. ‘‘scabrosa’’ Ukraine 1 CBFS JV6198 KC416121 C. ‘‘scabrosa’’ Ukraine 2 CBFS JV6199 KC416123 C. substerilis Austria CBFS JV7257 KC416107 C. substerilis Bulgaria CBFS (Exs. of Caloplaca, nr 11) KC416108 C. substerilis Czech Republic, holotype CBFS JV7920 KC416109 C. substerilis Slovakia PRA (Palice 13441) KC416110 C. ‘‘ulcerosa’’ USA GZU (Wetmore 93230) KC416105 C. ‘‘ulcerosa’’ USA 2 GZU (Advaita 4915) KC416106 C. sp. southern Russia CBFS JV8181 KC416141

Table 2. Summary of phylogenetic analyses: length of alignments (including gapped positions) and model selected for the purpose of MrBayes calculation

Target Group Phylogenetic tree Length of alignment Model

Caloplaca crenulatella group Fig. 2 525 positions SYM+ADGamma C. crenularia group Fig. 3 486 positions GTR+ADGamma C. xerica group Fig. 4 501 positions SYM+ADGamma Pyrenodesmia group Fig. 5 535 positions SYM+ADGamma C. ulcerosa and related taxa Fig. 6 519 positions GTR+ADGamma

Phylogenetic analyses ture in this paper, because generic names are still miss- ing for many Teloschistaceae taxa, including Caloplaca Five independent phylogenetic analyses of the nuclear emilii and C. substerilis described here. Names of other ITS region were made to cover the individual groups lichen taxa follow the Index Fungorum http://www. studied. All analyses followed almost the same design; indexfungorum.org/names/names.asp differences are listed in Table 2. Sequences were aligned using the MAFFT v6 server (http://mafft.cbrc.jp/alignment/server; Katoh & Toh 2008) according to the L- INS-i strategy. The resulting alignments required some Records new to Central Europe manual adjustments (done in BioEdit; Hall 1999) and, in the case of the C. crenularia group, also trimming of Caloplaca interfulgens (Nyl.) J. Steiner unalignable positions (using TrimAl-automated1 algo- 52: rithm, Capella-Gutierrez et al. 2009). The length of Verh. zool.-bot. Ges. Wien 479 (902). – Lecanora inter- 56: datasets submitted to further analyses ranged from 486– fulgens Nyl. Flora 340 1878. 535 positions. Final alignments were submitted to Tree- Images of some Czech and German specimens are avail- Base http://treebase.org/treebase-web/home.html. able on the lichenological web page at the University Molecular phylogenies were estimated by Bayesian in- of South Bohemia http://botanika.bf.jcu.cz/lichenology/ ference as incorporated in MrBayes 3.0b4 (Huelsenbeck index.php?pg=5. & Ronquist 2001; Ronquist & Huelsenbeck 2003). Model selection was committed to the Kakusan4 algo- Diagnostic characters. Thallus well-developed, rithm (Tanabe 2011), whereas the baseml software consisting of yellow areoles and often with (Adachi & Hasegawa 1996) served as the computational core. With reference to the Bayesian information criterion squamules at the margin. Ascospores polari- (Schwartz 1978), we opted for SYM or GTR models locular, c.15–19 50–75 mmwithseptaup with rate variation across sites simulated by discrete to 4 mm wide. Prothallus indistinct. Occurs on gamma distribution (G8) and autocorrelated by the calcareous rocks. AdGamma rates prior (Table 2.). The increased proba- Similar taxa are Caloplaca crenulatella s. lat. bility of transitions over transversions, well documented in many rDNA datasets (see e.g. Keller et al. 2007), was (the yellow thallus usually reduced), C. dif- reflected by setting the substitution rates prior (revMatPr) fusa Vondra´k & Llimona (on non-calcareous to dirichlet with values 1 and 3 for these two mutational rocks, yellow thallus with thin diffuse mar- types, respectively. Each analysis comprised two inde- gin, with grey-white prothallus, squamules pendent runs, each of which encompassed four Metro- polis-coupled MCMC chains with 10 000 000 genera- absent) and species of the Caloplaca velana tions sampled after every 1000th generation. In every complex (ascospores shorter with thicker run, one Markov chain was cold and three were incre- septa). mentally heated by the parameter of 03. To eliminate trees sampled before reaching apparent stationarity, the Distribution (Fig. 1A). Caloplaca interful- first 25% of entries were discarded as burn-in and the gens was previously known only from deserts, rest were used to compute majority-rule consensus, semi-deserts or steppes in North Africa where the relative occurrences of nodes are identified with the Bayesian posterior probabilities (Figs 2–6). (Nylander 1878; Navarro-Rosine´s & Hladun Bayesian posterior probabilities b50 are shown, branches 1996), Mediterranean Europe (Italy: Nimis with lower posterior probabilities are collapsed. & Martellos 2008; Spain: Nimis et al. 1998), Iran, Kazakhstan (Vondra´k et al. 2011) and Nomenclature continental Turkey (Vondra´k et al. 2012a). Arup et al. (2013) proposed a new nomenclature within The new records are surprisingly from less Teloschistaceae and split the crustose genus Caloplaca into arid territories in Austria, the Czech Repub- numerous genera. We do not follow the new nomencla- lic, Germany, southern Russia and Slovakia.

Fig. 1. Distribution maps. A, Caloplaca interfulgens, previously published data (white dots), new records (black dots); B, C. emilii (black dots), C. areolata (white dots).

In all Central European localities, C. interful- Taxonomic note. Although the Caloplaca gens is restricted to limestone outcrops in crenulatella group has been studied recently xerothermic sites (often south-facing steppes). (Navarro-Rosine´s & Hladun 1996; Vondra´k et al. 2011), it is still poorly understood and Phylogeny. In the ITS phylogeny of the many lineages are not yet well characterized. Caloplaca crenulatella group (Fig. 2), Calo- Fortunately, its well-developed areolate placa interfulgens forms a well-resolved sister thallus separates C. interfulgens from the many clade to Caloplaca tominii, a sorediate species taxa with reduced thalli. However, some with a similar distribution pattern in Europe. Central Asian taxa have a thallus similar to

Fig. 2. Bayesian ITS phylogeny of the Caloplaca crenulatella group; C. interfulgens clade delimited by the grey square.

C. interfulgens (e.g. ‘Caloplaca sp., southern outcrops 1 km NW of ruin of Karlı´k, 49560 5600N, Russia’ in Fig. 2) and their delimitation re- 14140 4900E, 300 m, 2011, I. Frolov & J. Vondra´k 9399 (CBFS); Praha, Radotı´n, Kosorˇ, protected area Cˇ erna´ quires further study. rokle, E of village, 49590 2100N, 14200 800E, 250–300 New records. Austria: Niedero¨sterreich: Wien, Hain- m, 2011, Z. Palice & J. Vondra´k 9144 (CBFS). Southern burg an der Donau, rocks on SW slopes of hill Braun- Moravia: Pavlovske´ vrchy hills, Mikulov, Klentnice, SE 0 00 0 00 berg NE of town, 48090 1000N, 16570 1200E, 280 m, slope of hill Pa´lava, 48 51 22 N, 16 38 33 E, 350–400 2012, J. Vondra´k 9550 (CBFS).—Czech Republic: m, 2012, J. Vondra´k 9577 (CBFS).—Germany: Baden- Central Bohemia: Bohemian karst, Beroun, Tma´nˇ, pro- Wu¨rttemberg: Schwa¨bishe Alb Mts, Langenau, Albeck, tected area Koty´z, 15 km NE of village, 49540 5600N, shallow valley W of Kornbergho¨fe, 520–550 m, 1984, 1420 5500E, 350 m, 2011, J. Vondra´k 9153, 9155 & V. Wirth 29418 (STU; hb. Wirth). Rheinland-Pfalz: Eifel 9156 (CBFS); Praha, Dobrˇichovice, Karlı´k, limestone Mts, Uxheim, Dreimu¨hlen, limestone outcrops in dry

grassland, 1992, V. Wirth 23937 &R.Du¨ll (STU; hb. Pycnidia not present on the available iso- Wirth).—Russia: Orenburgskaya Oblast’: surroundings type material. The type material is also de- of water reservoir ‘‘Iriklinskoe vodokhranilishche’’, vill. Chapaevka, limestone rocks on opposite slope of lake, scribed in Søchting et al. (2008). 0 00 0 00 NE of village, 52 05 12 N, 58 48 1 E, 270–290 m, Observations of the Central European speci- 2011, I. Frolov & J. Vondra´k 9396 (CBFS).—Slovakia: Cerova´ vrchovina upland: Filakovo, Hajnaćˇka, hill Ragacˇ, mens. (Fig. 7A). lime-rich outcrop of volcanic pyroclastics in open beech- Thallus rough and scabrous by blastidia, oak forest, 48130 2500N, 19590 600E, 500 m, 2012, J. densely covering the thallus surface. Blastidia Vondra´k 10137 (CBFS). Muranska´ Planina Mts: Brezno, (30–)58–67–72(–130) mm diam. [30; 3; 28]. Tisovec, hill Okruhla skala, c. 2 km W of town, 48400 4200N, 19540 5700E, c. 800 m, 2011, J. Vondra´k Thallus surface pale grey to white, but tips of 9260 (CBFS). Stra´zˇovske´ vrchy Mts: Ilava, Zliechov, on blastidia often dark grey. Grey pigmented S-slope of Mt Stra´zˇov, 48560 5900N, 18270 1600E, 1000 thallus parts containing Cinereorufa-green m, 2012, J. Vondra´k 10198 (CBFS). Vihorlat Mts: So- (green-grey in water, K-- , N+ red) in the brance, Podhorod [Podhradı´], 1930, J. Bucˇek (BRA, uppermost thallus cells. Thallus divided into sub Caloplaca zimmermannii, Servı´t, nomen ined.). thin and flat angular areoles, c.02–10mm Caloplaca scabrosa Søchting, Lorentsen diam. The real cortex absent, but indistinct & Arup alveolate cortex present in spots, of spherical, Nova Hedwigia 87: 89 (2008). thick-walled cells (walls c.1mm thick). Thallus without anthraquinones, but with atranorin. Images of European samples and the isotype are available on the lichenological web at the University of South Apothecia deep red (old blackened apo- Bohemia http://botanika.bf.jcu.cz/lichenology/index. thecia not observed), with anthraquinones; php?pg=5. major: parietin and 7-Cl emodin; traces of Observation of the type specimen. Isotype emodin, fragilin and parietinic acid (C+ pur- (CBFS JV9402, ex C; Søchting 5513) example owing to chlorinated compounds); bia- ined in detail. torine or zeorine; thalline exciple sometimes Thallus rough and scabrous by blastidia, strongly expanded in old apothecia. True ex- densely covering the thallus surface. Blastidia ciple of palisade prosoplectenchyma, of cells (40–)71(–130) mm diam. [10; 1; 29]. Thal- with glutinized, 1–2 mm thick walls. Inner lus surface pale grey to sordid white, but tips exciple and lower hypothecium brown-red of blastidia often dark grey. Grey thallus parts (perhaps by anthraquinones). Ascospores polar- containing Cinereorufa-green (green-grey in ilocular, (115–)130(–150) (65–)75 water, K-- , N+ red) in the uppermost thallus (–90) mm[10;1;14&08], with septa cells. Thallus divided into thin and more (30–)40(–50) mm[10;1;05]. or less flat angular areoles, c.02–13mm Pycnidia with red tops, containing chlori- diam. The real cortex absent, but indistinct nated anthraquinones (C+ purple). Conidia alveolate cortex present in spots, of spherical, more or less bacilliform, c.3–4 1 mm. thick-walled cells (wall c.1mm thick). Thallus Importance of particular characters. Calo- without anthraquinones, but with atranorin. placa scabrosa shares many characters with Apothecia biatorine, deep red (old apo- other related taxa from the C. crenularia thecia somewhat blackened), with anthra- group (as defined in Fig. 3), so their diagnos- quinones; major: parietin and 7-Cl-emodin; tic power is rather low. They include: 1) traces of emodin, 7-Cl-citreorosein, 7-Cl- presence of Cinereorufa-green in the thallus; emodinal and parietinic acid (C+ purple ow- 2) apothecia with chlorinated anthraquinones ing to chlorinated compounds). True exciple (C+ purple); 3) structure of the true exciple; of palisade prosoplectenchyma, of cells with 4) brownish pigment in lower hypothecium glutinized, c.1mm thick walls. Lower exciple and inner true exciple; 5) pycnidia with red and lower hypothecium brown-red (possibly caps. due to small amount of anthraquinones; with Some characters are specific for C. scab- weak K+ purple reaction). Ascospores polariloc- rosa: 1) presence and size of blastidia; 2) ular, (120–)140(–170) (55–)65(–80) presence of atranorin in the thallus. We have mm[10;1;13&07], with septa (40–) tested the diagnostic power of the presence 50(–55) mm[10;1;05].

Fig. 3. Bayesian ITS phylogeny of the Caloplaca crenularia group including the Central European sample of C. scabrosa and C. ‘‘scabrosa’’ from the Eastern Carpathians.

of atranorin. We analyzed thalli of various 2) thallus thickness; 3) extent of the thalline species of the C. crenularia group: Caloplaca exciple. Based on our observations of nu- ammiospila (Ach.) H. Olivier (CBFS merous samples of the C. crenularia group, JV10223), C. crenularia (With.) J. R. Laundon these characters were very variable both (CBFS JV4596; 5608; hb. Z. Palice 7837; Po- within and between specimens of a single land, Nowak’s exsiccate 203 in GZU; Sardinia, species, so the differences are of little taxo- 1986, Poelt in GZU), C. ferruginea (Huds.) nomic importance. Ascospore size and sep- Th. Fr. (CBFS JV7224; 7256), C. furfuracea tum width also differ between the type and H. Magn. (Ural, hb. I. Frolov), and C. hun- the Central European collections, but this garica H. Magn. (CBFS JV3081). Atranorin difference may be merely a consequence of was detected in only one sample of C. crenu- the low number of available specimens and laria from the basalt outcrops in the Karko- measurements. nosze Mountains, W Sudetes (Nowak, Lich. Phylogeny. The ITS sequence of the Cen- Polon. Exs. n. 203), indicating that this C. tral European specimen of Caloplaca scabrosa crenularia specimen does not belong in the is placed in the basal polytomy of the ITS main C. crenularia clade. phylogeny of the C. crenularia group (Fig. The type specimen of Caloplaca scabrosa 3). It is perhaps closely related to the arctic- differs from the Central European material alpine C. ammiospila or boreo-montane C. in the following characters: 1) size of areoles; furfuracea.

Taxonomic notes. The epixylic taxon Calo- Thallus forming irregular spots, brown- placa furfuracea is very similar to C. scabrosa. grey or pale to dark grey, to several cm wide; It likewise produces blastidia (isidia accord- often starting on other crustose lichens; of ing to Arup & A˚ kelius 2009) of the same tightly arranged, angular to rounded, flat to size; tips of blastidia are also usually dark slightly convex, areoles or squamules, (03–) grey due to the Cinereorufa-green content 06–09–11(–26) mm diam. [70;7;04]. in the alveolate cortex. With the exception of Thickness of thallus 100–500 mm. Medulla the ecology, the only reliable character dis- well-developed only in thick thalli, but up to tinguishing C. furfuracea from C. scabrosa is 400 mm thick; medullary tissue formed of the absence of atranorin. loose prosoplectenchyma; medullary hyphae We have collected samples of a granular to c.2–3mm wide with walls thickened up to 1 blastidiate lichen in the subalpine belt of the mm. Algal layer 50–140 mm thick; algal cells Eastern Carpathians (‘‘Caloplaca scabrosa’’ in globose, c.5–20mm diam. Cortex developed Fig. 3). These saxicolous specimens are very in patches, up to 30 mm thick, not gelatinous; similar to both C. furfuracea and C. scabrosa. sometimes only alveolate cortex present. They appear to be closer to C. furfuracea in Epinecral layer often present, up to about 10 the ITS phylogeny but they share chemistry mm thick. Cortex cells or alveolate cortex and ecology with C. scabrosa. cells spherical, thin-walled, about 4–6 mm New records. Czech Republic: Northern Moravia: diam. Blastidia simple, globose, dark grey, Ry´marˇov, Karlov, central part of Velky´ kotel corrie, always present, produced at margins of on phyllitic overhanging rock, 1330–1340 m, 2002, Z. areoles or squamules, rarely also on their Palice 7024 (PRA); ibid., 50030 2000N, 17140 E, 1250– upper surface, (20–)53–65–95(–210) mm 1300 m, 2004, J. Vondra´k 1907, 1908 & 1909 (CBFS). diam. [60; 6; 36]. Extracellular crystals of C. ‘‘scabrosa’’. Ukraine: Eastern Carpathians: Svido- vets Mts, at glacial lake at bottom of glacial cirque in N calcium salts not observed in any thallus slope, 48150 4100N, 24130 2200E, on sun-exposed base- part. Pruina absent. Prothallus indistinct or rich sandstone boulders close to water, c. 1300 m, absent. Thallus frequently affected by brown 2007, J. Vondra´k 6199 (CBFS). hyphomycetes resembling species of Intra- lichen. Apothecia present in c. 50% of samples col- New species lected; rare in northern populations; (03–) Caloplaca emilii Vondra´k, Khodos., 05–07–09(–14) mm diam. [40;4;02]; Cl. Roux & V. Wirth sp. nov. zeorine. Disc in shades of brown (orange in young apothecia); true exciple usually yellow MycoBank No: MB 803332 (contrasting with disc); thalline exciple in Thallus grey or brown-grey, non-pruinose, of more or shades of grey; pruina absent. Hymenium less flat areoles, with Sedifolia-grey and without anthra- colourless, without distinct gelatinous matrix quinones. Dark grey blastidia always present at margins of thallus units. Mature apothecia zeorine, usually with and without extracellular oil drops, c.70– brown disc and more or less yellow true exciple, Ce 110 mm high; epihymenium ochre to green- purple (with chlorinated anthraquinones). Ascospores yellow. Hypothecium colourless, rarely with broadly ellipsoid, less than 15 mm long, with thick septa. extracellular oil drops, more or less flat, c. Pycnidial tops dark grey. Conidia ellipsoid, not bacilli- 100–300 mm high, formed of cells variable form. Type: Bulgaria, Black Sea coast, Kavarna, limestone in shape; subhypothecial algal layer present cliffs on seashore 15 km NE of Kamen Brjag, (algal cells underlying entire hypothecium). 43270 587600N, 28330 550200E, on coastal limestone Exciple c. 70–110 mm wide, formed of true outcrop above supralittoral zone, 6 April 2007, J. exciple, c.30–60mm wide, and thalline ex- Vondra´k 6600 (CBFS—holotype; KHER—isotype). ITS sequence of the holotype: KC416101. ciple, c.10–70mm wide. Upper part of true exciple of thin-walled spherical cells c.4– Images of the German sample are available on the lichenological web page at the University of South Bohemia 6 3–4 mm. Lower part of palisade pro- http://botanika.bf.jcu.cz/lichenology/index.php?pg=5. soplectenchyma of thin-walled cells c.5– 12 15–20 mm. Thalline exciple without (Figs 1B, 4, 7B) cortex or with indistinct alveolate cortex.

Paraphyses 20–25 mm wide in lower part, is a similar lichen; it is very common in Cen- but widening gradually to (25–)30–35– tral Europe, but occurs mainly on non- 40(–50) mm[30;3;05] in upper part; calcareous rocks (orange, C-- apothecia, rarely branched and anastomosed. Asci clavate, without chlorinated anthraquinones, blasti- c.50–70 15–20 mm. Ascospores polarilocular, dia usually overgrowing most of thallus sur- (80–)120–125–135(–150) (50–)70– face). 75–80(–95) mm[50;5;15&09], septa Phylogeny. In the ITS phylogeny (Fig. 4), (40–)50–55–60(–75) mm[50;5;09]. Caloplaca emilii is definitely placed in the C. Ascospore length/breadth ratio: (10–)15– xerica group (sensu Vondra´k et al. 2012b). It 17–18(–22) [50;5;03]; septum width/ forms a well-circumscribed clade (PP 1 0), ascospore length ratio: (030–)040–045– ¼ sister to C. areolata. Both taxa form a well- 047(–060) [50;5;01]. Extracellular crystals of calcium salts absent from all apothe- supported monophyletic group (PP ¼ 099). cial parts. Ecology and distribution. Caloplaca emilii Pycnidia not common (observed in only occurs on sun-exposed, usually horizontal, three samples), c. 150–200 mm wide, with faces of limestone outcrops in fast-drying several partly separated chambers (Xantho- places in steppes, forest-steppes or in open ria-type), distinguished by their darker grey Mediterranean shrub vegetation, mainly in tops on the thallus surface. Conidiophores the Placocarpetum schaereri (Roux 1978: 120– formed of isodiametric cells, c.2–4mm 130). Co-occurring lichens are Acarospora cer- diam. Conidia ellipsoid, broadly ellipsoid or vina, Aspicilia calcarea, A. contorta, Bagliettoa tear-shaped, rather uniform in size, 20– calciseda, Caloplaca aurantia, C. chalybaea, C. 25 15 mm. coronata, C. crenulatella s. lat., C. inconnexa, C. lactea, C. teicholyta, C. variabilis, Cande- Chemistry. True exciple, medulla and lariella aurella, Diplotomma hedinii, D. venus- lower cortex non-amyloid (I-- ); hymenium tum s. str., Heteroplacidium fusculum, Lecanora and hypothecium amyloid (I+). Uppermost muralis s. lat., Lobothallia cheresina s. lat.,L. cells in cortical tissue of thallus and thalline radiosa, Placocarpus schaereri, Placopyrenium exciple contain Sedifolia-grey (grey in water, canellum, Rinodina calcarea, R. ocellata, R. K+ violet, N+ red, H2SO4+ red, I+ blue). Content of Sedifolia-grey is higher in pycni- bischoffii, Verrucaria lecideoides, V. macrostoma f. furfuracea,andV. nigrescens s. lat. dial tops. Epihymenium and outer cells in The species is already known from Ger- the true exciple contain anthraquinones: fragilin (major) and 7-Cl-emodin (HPLC done many (as the blastidiate variant of Caloplaca areolata in Wirth et al. 2011). Nevertheless, in sample JV6597). this lichen has a rather southern distribu- Etymology. The epithet is derived from the tion in Europe; it is probably most common name of our great friend Emil Cˇ ervenka, who in the Mediterranean basin and adjacent supported the first author during difficult areas, such as France, Italy, Spain, mainly in times. the supramediterranean and montane belts (Roux 1978: 124, as C. areolata). Although Similar taxa. Caloplaca areolata (Zahlbr.) it is common in continental areas around Clauzade (without blastidia), C. chlorina the Black Sea, we do not know it from conti- (Flot.) Sandst. and C. isidiigera Veˇzda (with nental areas east of the Mediterranean basin. blastidia but with lecanorine apothecia and In southern areas, it sometimes grows with bacilliform conidia), C. concreticola Vondra´k its close relative C. areolata (for example in & Khodos. (with blastidia but without anthra- southern France and Greece). Both taxa quinones in apothecia), C. soralifera Vondra´k have similar ecology, but C. areolata without & Hrouzek (with soredia, often pruinose) vegetative diaspores appears to be restricted ˇ and C. xerica Poelt & Vezda (usually with to the Mediterranean region, whereas the isidia, without flat areoles, with larger asco- blastidiate C. emilii also occurs in isolated spores). A little-known blastidiate morpho- localities far to the north (Fig. 1B). The type of Caloplaca atroflava (Turner) Mong.

C. demissa AF353961 0.97 C. haematites JN641769 C. aractina GU723413 C. tianshanensis GU552276

0.95 C. areolata Spain 1 C. areolata Montenegro C. areolata Croatia 0.99 0.96 C. emilii Bulgaria, Rhodopes C. emilii France 1 C. emilii Bulgaria HOLOTYPE C. emilii Czech Republic 1

0.93 C. emiliii Czech Republic 2 C. emilii Greece C. xerica JN641792 0.71 0.96 C. atroflava HQ234606 1 C. atroflava HQ611276 0.75 C. soralifera JN641780 1 C. teicholyta EU639593 C. erythrocarpa EU639591 0.97 0.78 C. fuscoatroides JN813404 1 C. fuscoatroides HQ234599 . 0 06 0.95 C. ceracea HQ234600 C. ceracea HQ234603

Fig. 4. Bayesian ITS phylogeny of the Caloplaca xerica group including C. emilii (in the grey square) and C. areolata.

ability to reproduce vegetatively may have also under ecology) and C. isidiigera is an facilitated the northward extension of its unrelated species with lecanorine apothecia distribution. A similar situation is observed and a (sub-)alpine distribution (Veˇzda 1978; in other Mediterranean lichens from the C. Sˇ oun et al. 2011). xerica group; sorediate/blastidiate C. albo- Caloplaca areolata has recently been con- lutescens (Nyl.) H. Olivier and C. teicholyta sidered a synonym of C. spalatensis Zahlbr. (Ach.) J. Steiner are known from much (e.g. Nimis & Martellos 2008). This syno- more northern territories than the closely re- nymization is incorrect, because C. spalaten- lated C. erythrocarpa (Pers.) Zwackh, which sis is a very different lichen which belongs to is without vegetative diaspores. the C. crenularia group (images of both holo- types, deposited in the herbarium W, are Taxonomic notes. Caloplaca emilii is well available on the lichenological web page at known from the Mediterranean regions of the University of South Bohemia http://bota- France, where it has been named C. areolata nika.bf.jcu.cz/lichenology/index.php?pg=5). (Clauzade 1963, 1965, 1969; Roux 1978) or later C. isidiigera (Roux 1982, 1984; Bois- Paratypes. Austria: Niedero¨sterreich: Wien, Hainburg sie`re et al. 1989; Houmeau & Roux 1991; an der Donau, rocks on SW slopes of hill Braunberg NE of town, 48090 1000N, 16570 1200E, 280 m, 2012, Roux & Gueidan 2002; Bricaud 2007). J. Vondra´k 9570 (CBFS).—Bulgaria: Black Sea coast: However, these names belong to other taxa; Kavarna, Kamen Brjag, 43270 5900N, 28330 5500E, C. areolata lacks vegetative diaspores (see 2007, J. Vondra´k 6600 (CBFS). The Rhodopes: Mad- zharovo, Silen, Byal Kladenets, in valley below village,

41370 N, 25400 E, 350 m, 2004, J. Vondra´k 2223 Thallus epilithic, ochre, white-grey to dark (CBFS).—Czech Republic: Southern Moravia: Miku- grey, usually with white pruinose spots, lov, in town, ruin of castle Kozı´ Hra´dek, 48480 3400N, 16380 1700E, 2011, J. Vondra´k 9358 & O. Vondra´kova´ forming irregular spots to several cm wide; (CBFS); Mikulov, Klentnice, SE slope of hill Pa´lava, of tightly arranged, angular to rounded, 48510 2200N, 16380 3300E, 350–400 m, 2012, J. Von- more or less flat areoles or somewhat um- dra´k 9581 (CBFS); Mikulov, Klentnice, at ruin of bilicate squamules, (0 44–)0 70–0 95–1 26 0 00 0 00 Sirotcˇ´ı hra´dek, 48 50 43 N, 16 38 25 E, c. 410 m, (–2 05) mm diam. [100; 10; 0 35]. Marginal 2011, J. Vondra´k 9357 & O. Vondra´kova´ (CBFS).— France: Provence: Vaucluse, Gordes, entre les Devens areoles sometimes bigger than areoles in the et Lancie, sur dalle de molasse mioce`ne au ras su sol, centre. Several small, tightly arranged areoles 439026N, 51931E, 275 m, 1975, G. Clauzade may merge to form larger units, but on the (MARSSJ 189); Vaucluse, Mirabeau, 520 m, 2005, C. contrary, large areoles are sometimes divided Roux 23475 (hb. Roux).—Germany: Bayern: Ober- franken, Fra¨nkische Alb: Kleinziegenfelder Tal, Grenz- into smaller subareoles due to secondary stein, 1976, V. Wirth 6101 (STU).—Greece: Attica: crevices. Thickness of the thallus, together Poros, limestone outcrops in N-part of island, with brown (probably necrotic) lower me- 37310 2800N, 23290 1000E, c. 200 m, 2010, J. Vondra´k dulla (02–)06–12–22(–50) mm [30; 3; 8726, 8832 & O. Vondra´kova´ (CBFS).—Romania: 10]; thickness of the thallus without lower- Dobrogea:Taˆrgus¸or, 44270 462600N, 28280 075900E, 2007, J. Vondra´k 6599 (CBFS); Tulcea, Enisala, most brown part (01–)03–04–05(–09) 44520 420900N, 28510 012700E, 2007, J. Vondra´k mm [30;3;02]. The brown lower medulla 6604 (CBFS); Tulcea, Popina Island, 44580 0300N, usually distinct, up to 125 times thicker 28580 5700E, 2007, J. Vondra´k 6596, 6597, 6598 & than the rest of the thallus. Colourless me- 7149 (CBFS).—Ukraine: Kherson region: Berislav, Bur- gunka, 2008, A. Khodosovtsev & G. Naumovich (KHER, dulla also present, (50–)140–235–330(–550) dupl. in CBFS). mmthick[26; 3; 145]; cells hardly observable due to presence of extracellular crystals in- Caloplaca molariformis Frolov, soluble in KOH and only partly dissolved Vondra´k, Nadyeina & Khodos. sp. nov. and recrystallized into needles in H2SO4. Algal cells arranged in vertical stacks, (30–) MycoBank No: MB 803333 67–91–129(–250) wide [47; 6; 44], and Anthraquinones entirely absent. Thallus epilithic, thick, (100–)223–263–334(–550) mm high [47;6; ochre or dark grey, pruinose in spots, with Sedifolia-grey 112]. Algal cells globose, (8 0–)12 6–13 7– in superficial fungal cells. Blastidia and/or soralia always present. Thallus formed by high algal and fungal stacks 145(–220) mm diam. [30;3;32]. Cortex (sensu Vondra´k & Kuba´sek 2013). Fungal stacks of col- above the algal stacks absent or indistinct, ourless palisade prosoplectenchyma, of cells elongated alveolate cortex present, up to c.15–30mm vertically. The upper thallus surface with ridges derived thick; upper fungal cells in algal stacks grey, from the epinecral layer, above fungal stacks (similar structure is described in South African ‘‘Fensterflechten’’ containing Sedifolia-grey. Fungal stacks (mea- by Vogel 1955). Epihymenium and outer part of true sured with epinecral layer) (13–)45–86– exciple brown to grey, with Sedifolia-grey, K+ (slightly) 120(–270) wide [46; 6; 55] and (75–)180– violet to violet-brown. Ascospores c.14–18mm long 322–505(–750) mm high [46; 6; 165]; with rather thin septa, c.3mm wide. formed by vertically oriented palisade proso- Type: Slovakia, Cerova´ vrchovina upland, Filakovo, Hajnaćˇka, Sˇ urice, SW-slope of the hill Sovı´ hrad, plectenchyma; size of cells in the middle part 48130 3400N, 19540 4500E, on lime-rich outcrop of vol- of stacks (45–)94–119–133(–180) canic pyroclastics in sun-exposed abandoned quarry, c. (30–)37–43–48(65) mm[30;3;39& 250 m, 8 November 2012, J. Vondra´k 10192 (CBFS— 09]. In lower part of stacks, cells longer and holotype; isotypes to be distributed in Exsiccates of Caloplaca, fasc. 4). ITS sequence of the holotype: narrower; in uppermost part, cells almost KC416142. isodiametric, c.4–7mm diam. Epinecral layer More images available on the lichenological web page above fungal stacks usually well-developed, at the University of South Bohemia http://botanika.bf. (5–)20–95–200(–350) mm thick [81; 9; 72]; jcu.cz/lichenology/index.php?pg=5. dead cells (colourless in cotton blue) recognizable in the lower part. Boundary between (Figs 5, 7C; fig. 2 in Vondra´k & Kuba´sek epinecral layer and upper cells of the fungal 2013) stack sometimes indistinct, but recognizable after KOH treatment as a sordid grey-violet

Fig. 5. Bayesian ITS phylogeny of the Pyrenodesmia subgroup of Caloplaca, including C. molariformis clade delimited by the grey square.

line caused by traces of the Sedifolia-grey in (always present) or rarely soredia; sometimes uppermost fungal stack cells. Epinecral layer diaspores poorly developed, present only on often forms distinct ridges on thallus surface few areoles. Blastidia simple, more or less above fungal stacks, because it is absent from globose, (30–)54–67–89(–150) mm diam. surface of algal stacks (Fig. 7C). Epinecral [52; 6; 25], dark grey, present on the margin ridges best developed in samples from de- and upper surface of areoles and squamules; serts of Western Kazakhstan, but less distinct detached blastidia occasionally cover the in samples from Slovakia and Ukraine. Fun- whole surface. Blastidia sometimes with ap- gal stacks sometimes reaching medulla at the pearance of consoredia, with internal sore- bottom and the boundary between the stacks dia-like structures. Extracellular crystals sol- and medulla recognized by the crystals abun- uble in KOH and Sedifolia-grey pigment dant in medulla but absent from stacks. Mar- present in outer fungal cells of blastidia. gins of areoles and squamules and the lower Soralia rarely observed, on the upper surface surface of squamules usually with cortex, up between epinecral ridges; soredia c.25–40 to c.20mm thick, of isodiametric cells, c.4–7 mm diam. White pruina always present, mm diam. Vegetative diaspores are blastidia better developed between epinecral ridges.

Prothallus indistinct or absent. Thallus fre- mm[30; 3; 1 & 10]. Ascospores polarilocular, quently affected by brown hyphomycetes re- (120–)143–162–183(–230) (50–) sembling species of Intralichen. 64–77–91(105) mm[70;8;23&13]; Apothecia (033–)042–055–072(–132) septa (20–)26–30–33(–40) mm wide mm diam. [100; 10; 015], zeorine or rarely [70;8;05]. Ascospore length/breadth ratio: almost lecanorine; mature apothecia sessile, (140–)198–212–227(–286) [70;8;032]; usually not abundant on thallus, sometimes septum width/ascospore length ratio: (011–) absent. Richly fertile populations known 017–019–022(–030) [70;8;004]. Asco- only from Slovakia and Ukraine. Disc brown spores with well-developed septa often absent. to black, not pruinose, sometimes cracked; Pycnidia rare, c. 140–190 mm wide, mainly true exciple concolourous with the disc, oc- with a single chamber, present on the upper casionally white pruinose; thalline exciple thallus surface, but also on the lower surface concolourous with the thallus, with white of squamules; superficially hardly distinguish- pruina. Hymenium (63–)91–102–109(–175) able. Old pycnidial chambers sometimes mm high [30; 3; 23], colourless, often with filled by crystals insoluble in KOH. Coni- very small (<1 mm) extracellular oil drops, diophores of spherical or triangular, more or sometimes strongly inspersed with extra- less isodiametric cells. Conidia narrowly to cellular oil drops up to c.2mm diam., broadly ellipsoid, 25–45 15–20 mm sometimes not inspersed; without crystals. [14;2;02&05]. Epihymenium brown, grey or grey-brown. Chemistry. Spot tests: thallus Ke violet Hypothecium colourless, underlain by the (sometimes not observable or observable only algal layer, usually with extracellular oil in spots with blastidia and soredia), apothecia drops, without extracellular crystals; with a K-- , thallus and apothecia C-- ,P-- ,UV-- . central conical extension downward, (75–) Epihymenium, uppermost true exciple, upper- 153–174–185 (–275) mm high [30; 3; 48]; most fungal cells in thallus and vegetative formed of thin-walled cells variable in shape. diaspores contain Sedifolia-grey (grey or in- Exciple c. 10–160 mm wide. True exciple (10–) visible in water, K+ sordid violet). The re- 18–35–54(–93) mm wide [30; 3; 22], and action above fungal stacks usually weaker thalline exciple (0–)18–24–27(–68) mm than above algal stacks. Strongest reaction wide [30; 3; 20]. Upper part of the true ex- in superficial hyphae of vegetative diaspores. ciple grey-brown, brown-grey or grey, of True exciple non-amyloid (I-- ); hymenium thin-walled cells (40–)62–66–73(–100) and hypothecium amyloid (I+). No substan- (20–)34–45–52(–80) mm[100; 10; 11 ces revealed by HPLC (apothecia and thallus &11]. Lower part colourless, of palisade of an isotype were investigated). prosoplectenchyma of thin-walled cells (60–) 77–82–87(–115) (20–)24–28–33 Etymology. Areoles and squamules of the (–50) mm[30;3;13&08]. Thalline exciple lichen thallus often resemble molars of her- sometimes with cortex in its upper part, c.8– bivores. 20 mm thick; cortex changing into alveolate cortex in the lower part of thalline exciple. Similar taxa. The thallus anatomy, with Cells of the cortex spherical, c.35–70 mm tissues in stacks, is very rare within the Pyre- diam., often hardly observed due to extracel- nodesmia subgroup of Caloplaca. It is present lular crystals insoluble in KOH. Paraphyses in one known species only, Caloplaca albo- (15–)21–23–28(–35) mmwide[100; 10; variegata (B. de Lesd.) Wetmore, which is 04] in lower part, but widening gradually to very similar to C. molariformis but has no vege- (30–)35–44–55(–65) mm[100; 10; 08] tative diaspores (Wetmore 1994; lectotype in in upper part; rarely branched and anas- UPS seen). This species was described from tomosed; the uppermost cell of paraphyses North America, but similar morphotypes are usually dead and deformed. Asci clavate, known in continental Eurasia (our observa- (40–)58–64–69(–85) (12–)17–20–21(–28) tions). Zhou et al. (2012) reported a taxon with tissues in stacks from China and named

it C. albovariegata, but it has a thallus surface Beyneu, c. 50 km SW of town at road to Aktau, valley of without ridges derived from the epinecral salt river Manashi, 45010 2600N, 54590 5600E, 50 m, 2009, A. Khodosovtsev & J. Vondra´k 9483 (CBFS); layer and it does not resemble C. molarifor- Mangistau district, West Aktau ridge, soft valley with mis. Other similar taxa are Caloplaca albopus- rocky outcrops at river Akespe, 44240 2100N, 51350 5900E, tulata Khodos. & S.Y. Kondr. (with pustules 100 m, 2009, A. Khodosovtsev & J. Vondra´k 9486 and schisidia), C. bullata (Mu¨ll. Arg.) Zahlbr. (CBFS); Mangistau district, at road between village (bullate thallus without vegetative diaspores), Shetpe and Say-Utes, c. 30 km SW of Say-Utes, 44090 2000N, 52390 1000E, 260 m, 2009, A. Khodo- C. concreticola (with soralia) and C. trans- sovtsev & J. Vondra´k 9506 (CBFS); Mangistau district, caspica (Nyl.) Zahlbr. (without vegetative East Karatau ridge, rocks at road between Zhatybay diaspores), but all these taxa have thallus and Shetpe, c. 30 km SW of Shetpe, 43570 0000N, tissues arranged in horizontal layers, not in 52050 5200E, 180 m, 2009, A. Khodosovtsev & J. Von- dra´k 9499 (CBFS).—Russia: Orenburgskaya Oblast’: stacks. They also do not have specific ridges Orenburg, village Mikhaylovka (c. 30 km SES of city), derived from the epinecral layer. (Type Khanskaya gora hill, S of village, above brook Ber- specimens and other comparative material d’yanka, 51250 4800N, 55260 2700E, c. 200 m, 2011, I. studied by the authors.) Frolov & J. Vondra´k 9456 (CBFS); Saraktash district, protected area Kamennaya, rock outcrops in S-slope Phylogeny. In the ITS phylogeny (Fig. 5), above river Sakmara, 51560 5300N, 55580 2300E, 180 m, Caloplaca molariformis is placed in the C. 2012, I. Frolov & J. Vondra´k 10225 (CBFS). Republic of Altay: Kosh-Agach district, Kosh-Agach, Telengit- variabilis group, closely related to C. albopus- Sortogoy, S-slopes of Kuray Ridge (easternmost part), tulata. c. 6 km N of village, 50040 2400N, 88420 3000E, 2000– 2100 m, 2012, I. Frolov & J. Vondra´k 10224 Distribution and ecology. Caloplaca molari- (CBFS).—Slovakia: Cerova´ vrchovina upland: Fila- formis is mainly distributed in steppes and kovo, Hajnaćˇka, Sˇ urice, SW-foot of hill Sovı´ hrad, deserts of Iran, Kazakhstan, continental Tur- 48130 3400N, 19540 4500E, 240–250 m, 2012, Z. Fakov- key and southern Russia, at altitudes of 50– cova´, A. Guttova´, J. Lisˇka, Z. Palice 15905 & J. Vondra´k 10190 (CBFS, PRA; topotypes).—Turkey: Eastern 2100 m. Two isolated localities are also Anatolia:Ig˘dır, shale hills SE of town, 39510 2300N, known from the steppe or forest-steppe, in 44050 4200E, 1060 m, 2007, J. Vondra´k 6463 (CBFS). eastern Ukraine and southern Slovakia. The Central Anatolia: Yozgat, Bog˘azlıyan, O¨ zler village, species occurs in sunny habitats on soft lime- 39040 1000N, 35080 1700E, 1100 m, 2012, J. Vondra´k stone, chalk, calcareous sandstone or tuffs 9751 (CBFS); Kayseri, Talas, Derevenk valley, 38410 2300N, 35340 5200E, 1230 m, 2012, J. Vondra´k with evident content of lime (always reacting 9760, 9809 & 9787 (CBFS); Kayseri, south-east of with HCl). Co-occurring lichen taxa include Himmetdede, north-west of Kalkancık village, montane Acarospora spp., Aspicilia spp., Caloplaca steppe with shrubs, 38530 4300N, 35070 0100E, 1170 m, concreticola, C. crenulatella s. lat., C. decipiens, 2012, J. Vondra´k 9791 (CBFS).—Ukraine: Donetsk Upland: Luhansk region, Lutugyno district, steppe C. flavocitrina, C. soralifera, C. sororicida, C. slopes with marl outcrops near village Rozkishne, in teicholyta, C. tominii, C. transcaspica s. lat., botanical reserve ‘‘Balka Ploska’’, c. 150 m, 2007, O. C. xerica, Candelariella aurella, Lecanora Nadyeina 131, 132 & 134 (KW). [Specimens from muralis s. lat., Lemmopsis arnoldiana, Lichi- Ukraine were published as Caloplaca concreticola in nella sp., Verrucariaceae spp. (e.g. Staurothele Nadyeina (2009)]. frustulenta, Verrucaria macrostoma, V. nigrescens agg.). Caloplaca substerilis Vondra´k, Palice & van den Boom sp. nov. Paratypes. Iran: West Azerbaijan: Lake Urmia, rocks at road c. 2 km N of Saraydeh, 37520 5900N, MycoBank No: MB 803334 0 00 45 34 26 E, 1280 m, 2007, J. Vondra´k 5556 (CBFS); Similar to Caloplaca ulcerosa, but differs in thallus mor- 0 00 0 00 Khoy, airport, 38 25 1617 N, 44 54 2405 E, 1180 phology. Thallus endophloeodal, but also forming minute m, 2007, J. Vondra´k 5801 (CBFS); Lake Urmia, rocky areoles or squamules; sorediate; without any pigments 0 00 outcrops near coast N of Aq Gonbad, 37 49 1202 N, or TLC identifiable compounds. Apothecia up to c.05 0 00 45 25 0961 E, c. 1290 m, 2007, J. Vondra´k 5846 mm diam., orange-red, not pruinose, without chlori- (CBFS).—Kazakhstan: Mangistau province: Mangistau nated anthraquinones, biatorine to zeorine. Ascospores district, village Shetpe, West Karatau ridge, c.15kmN broadly ellipsoid, c.10–15mm long, with septa c.4–6 0 00 0 00 of village, 44 14 35 N, 52 03 19 E, 100 m, 2009, A. mm wide. Pycnidia with yellow caps containing anthra- Khodosovtsev 7775–7781 & J. Vondra´k 8262, 8247, quinones. Conidia bacilliform, c.3–4 10–15 mm. 9477 & 9487 (CBFS, KHER); Beyneu district, village

Fig. 6. Bayesian ITS phylogeny of Caloplaca ulcerosa and related taxa including C. substerilis (delimited by the grey square).

Type: Czech Republic, Southern Bohemia, Novoh- large areas of trunks. Squamules 100–150 mm ˇ ˇ radske´ hory Mts, Benesˇov nad Cernou, Zofı´n, alt. 745 thick and (010–)017–018–019(–030) mm m, 48400 2900N, 14410 3800E, on bark of solitary Ulmus glabra, 26 May 2010, J. Vondra´k 7920, A. Vondra´kova´ diam. [30;3;005]. Soralia small, usually & O. Redchenko (CBFS—holotype). ITS sequence of extended in one direction (rarely rounded), the holotype: KC416109. usually up to 02 mm in length, formed in More images available on the lichenological web page at tiny cracks in the tree bark or on margins the University of South Bohemia http://botanika.bf. and lower surface of squamules, usually not jcu.cz/lichenology/index.php?pg=5. in concave, crater-like depressions (typical (Figs 6, 7D) for Caloplaca ulcerosa Coppins & P. James); soralia in older lichens often tightly arranged Thallus endophloeodal or partly of diffuse and may resemble a continuous sorediate tiny squamules (somewhat epiphloedal areo- crust. Soredia without pigmentation, (15–) late thallus present in samples from the 23–24–26(–30) mm diam. [40; 4; 4]; consor- Alps); sorediate; forming irregular pale grey edia (30–)37–41–46(–65) mm diam. [40;4; to white spots or extensive crusts, covering 8]. Fungal cells in soredia or consoredia

Downloaded from https:/www.cambridge.org/core. University of Basel Library, on 11 Jul 2017 at 09:02:28, subject to the Cambridge Core terms of use, available at https:/www.cambridge.org/core/terms. https://doi.org/10.1017/S0024282913000455 at Downloaded from https:/www.cambridge.org/core/terms 03New 2013 https:/www.cambridge.org/core . https://doi.org/10.1017/S0024282913000455 Teloschistaceae . University of BaselLibrary nCnrlEurope— Central in , on 11 Jul2017 at09:02:28 Vondra , subjectto theCambridgeCore termsofuse,available ´ k ta.717 al. et

Fig.7.A,Caloplaca scabrosa (CBFS JV1908); B, C. emilii (holotypus); C, C. molariformis, morphotype with well-developed epinecral ridges above fungal stacks (CBFS JV9486); D, C. substerilis (CBFS JV7920, holotypus). Scales: A & C ¼ 1 mm; B & D ¼ 05mm. 718 THE LICHENOLOGIST Vol. 45

(35–)54–55–57(–75) (20–)32–33– c.50–60 10–16 mm. Ascospores polarilocu- 34(–45) [20;2;11&08]. Surface of sore- lar, (100–)120–120–125(–165)(50–) dia papillate; papillae formed of fungal cell 75–80–80(–105) mm[20;2;14&13], outgrowths, up to 7 mm high. Medulla indis- septa (40–)45–50–55(–85) mm wide tinct or absent. Algal layer forms majority of [20; 2;13]. Ascospore length/breadth ratio: thallus, c. 100–140 mm thick; algal cells glo- (12–)16–16–17(–22) [20;2;03]; sep- bose, c.5–20mm diam.; old cells often inter- tum width/ascospore length ratio: (026–) nally divided into several irregularly spherical 034–040–042(–052) [20;2;01]. Extra- autospores (cell division typical for Trebouxia; cellular crystals of calcium salts absent from e.g. Peksa & Sˇ kaloud 2008). True cortex ab- all apothecial parts. sent; alveolate cortex developed in patches, Pycnidia more common than apothecia up to 20 mm thick, of thin-walled, more or (observed in c. 50% of samples), c. 50–100 less spherical cells. Epinecral layer indistinct. mm wide, with several partly separated cham- Thallus surface papillate; papillae of the bers (Xanthoria-type), distinguished by their same size and character as those in soredia. yellow tops containing anthraquinones. Con- Extracellular crystals of calcium salts not idiophores various in height, formed of rectan- observed in any thallus part. Pruina absent. gular, triangular or spherical cells, c.3– Prothallus indistinct or absent. 5 4–8 mm. Conidia usually bacilliform, Apothecia present in c. 20% of samples straight or slightly curved, rarely ellipsoid collected (indicated by asterisk in the list or tear-shaped, (25–)32–34–35(–50) of paratypes), but fertile specimens usually (10–) 12–12–13(–15) mm[20;2;07& with scattered apothecia. The sample from 02]. the Alps (van den Boom 15927) with many Chemistry. Spot tests: thallus K-- ,C-- ,P-- , apothecia is exceptional. Apothecia mostly UVe white; apothecia K+ purple, C-- ,UV-- . up to 05 mm diam.; biatorine or zeorine. True exciple non-amyloid (I-- ). Hymenium Disc orange to orange-red; true exciple yellow- and the upper part of hypothecium (sub- orange to orange (usually somewhat paler than -- disc); thalline exciple (when visible) yellow hymenium) amyloid (I+). The C reaction of epihymenium and outer cells in the true to white; pruina absent or indistinct. Hyme- exciple suggests an absence of chlorinated nium colourless, somewhat gelatinous, with- anthraquinones. No compounds revealed out extracellular oil drops, c.60–70mm from thallus by TLC. high; epihymenium ochre. Hypothecium colourless, up to 100 mm high, more or less ﬂat, Etymology. ‘Substerilis’ reﬂects the usually but with downward extension through the sterile occurrence. subhypothecial algal layer in the centre, of thin-walled cells variable in shape; extracel- Similar taxa. Apothecial characters in the lular oil drops not seen. Exciple c.40–80mm new species are identical to those of the wide, formed of true exciple, c.30–70mm closely related Caloplaca ulcerosa, but they wide, and thalline exciple, c.0–30mm wide. differ in thallus characters. In C. substerilis, Upper part of true exciple of cells c.4– the thallus is endophloeodal or of diffuse min- 8 3–5 mm, with thin or more than 1 mm ute squamules, with marginal soralia, while thick, glutinized walls. Lower part of palisade Caloplaca ulcerosa forms an epiphloedal non- prosoplectenchyma of thin-walled cells, 6– squamulose thallus with round to irregular 11 2–4 mm. Thalline exciple sometimes soralia formed in crater-like depressions. with alveolate cortex in lower part, up to 30 The latter species further differs in its mm thick, of spherical cells; thalline exciple shorter, ellipsoid conidia (c.25–30 15 sometimes sorediate. Paraphyses 15–20 mm mm), much higher fertility and in ecology; it wide in lower part, but about three upper cells is a maritime species (Vondra´k et al. 2009b). widened; branching and anastomosing not White morphotypes of C. phlogina (Ach.) observed; paraphyses tips (35–) 46–46– Flagey are similar (see Kondratyuk et al. 46(–55) mm[20;2;07] wide. Asci clavate, 1998; Vondra´k et al. 2010b); they also have

papillate soredia of similar size without pig- in the Southern Ural Mountains (most of mentation, yellow pycnidial caps and an en- known localities). It is probably distributed dophloedal thallus, sometimes with minute throughout eastern and central Europe in white squamules. However, C. phlogina dif- suitable woodland areas with preserved un- fers in frequently having apothecia: these are disturbed solitary elm and poplar trees. So large (mostly >05 mm diameter), yellow- far it is known from Austria, Bulgaria, the orange, with a rough surface caused by Czech Republic, Russia and Slovakia. yellow anthraquinone pruina. Ascospores Taxonomic notes. The North American are significantly smaller with thinner septa: taxon called Caloplaca ulcerosa (Wetmore ascospores (85–)105–108–112(–130) 2004) is morphologically more similar to C. (40–)50–54–57(–70) mm[30;3;10& substerilis than to C. ulcerosa s. str. We have 07], and septa (25–)31–36–38(–45) mm examined three samples of the North Ameri- wide [30;3;06]. can taxon (GZU: Iowa, Teloschistaceae Exsic- Sterile thalli may resemble a number of taxa, including: Caloplaca obscurella (J.Lahm) cati 95; Iowa, Wetmore 93230; South Dakota, Advaita 6490), and did not find any diag- Th. Fr. (with rounded crater-like soralia, nostic difference from C. substerilis. It corre- brown apothecia), C. sterilis Sˇ oun et al. (on sponds well with the ITS phylogeny, where steppe shrubs, with lecanorine apothecia), both taxa form a monophyletic group. The C. subalpina Vondra´k et al. (saxicolous, with distribution of the North American taxon lecanorine apothecia and Sedifolia-grey in (Wetmore 2009) and the distribution of C. soredia), Candelariella subdeflexa (Nyl.) Lettau substerilis are similarly continental and differ- (with different apothecia and more conspicu- ent from the maritime distribution pattern of ous squamules producing conidia from the C. ulcerosa s. str. (Vondra´k et al. 2009b). Pro- underside) and Rinodina degeliana Coppins (areolate-squamulose thallus with marginal visionally, we call the North American specimens C. ‘‘ulcerosa’’ in Fig. 6. soralia; presence of atranorin and zeorin). Two ITS sequences of C. substerilis from Phylogeny. In the ITS phylogeny (Fig. 6), the Alps form a separate lineage from the Caloplaca substerilis forms a well-supported other C. substerilis sequences. The specimens clade, sister to the clade of North American from the Alps also differ slightly in mor- C. ‘‘ulcerosa’’. Both taxa are sister to the Eu- phology (frequent apothecia, more or less ropean C. ulcerosa. Close relatives of these epiphloeodal thallus and absence of minute three taxa are not known. squamules) and ecology. While most samples were collected from solitary elms, pop- Ecology and distribution. Caloplaca substerilis lars and oak, specimens from the Alps came occurs on nutrient-rich bark of Acer campest- from Sambucus bark and spruce twigs. This ris, A. platanoides, Carpinus, Juglans, Quercus, suggests that the populations from the Alps Populus and Ulmus in well-lit conditions, might represent a distinct infraspecific taxon. sometimes overgrowing mosses on bark. Specimens from the Alps were collected on Paratypes (fertile specimens indicated by asterisk). Aus- the bark of Sambucus and on Picea abies tria: Steiermark: Schladming, Ramsau am Dachstein, in gorge with road from Ramsau to Weissenbach, c. 850 m, twigs. Co-occurring lichens are more or less 2009, J. Vondra´k 7257 (CBFS). *Ka¨rnten: Gailtaler nitrophilous Caloplaca cerinelloides, C. mona- Alpen, 10 km WNW of Weissbriach, 05 km SE of Fel- censis, C. obscurella, Lecanora hagenii, Macen- stritz, open pine forest, 550 m, 1994, P. van den Boom tina dictyospora, Phaeophyscia nigricans, P. 15927 (hb. van den Boom).—Bulgaria: The Rhodopes: Madzharovo district, Silen, Rabovo, valley of small orbicularis, Physcia spp., Physconia sp., Picco- brook N of village, 41370 N, 25400 E, 250 m, 2004, lia ochrophora, Rinodina pityrea, Xanthomen- J. Vondra´k (CBFS, in sample ‘‘Caloplaca virescens, Exs. doza fulva and Xanthoria parietina. of Caloplaca, Nr 11’’).—Czech Republic: Southern Caloplaca substerilis shows continental bias Bohemia: Novohradske´ hory Mts, Benesˇov nad Cˇ ernou, ˇ 0 00 0 00 in Europe. It appears to be quite common Zofıń, 48 40 29 N, 14 41 38 E, 745 m, 2009, 2010, Z. Palice 12943 & 13676 (PRA, topotypes); Sˇ umava Mts, Borova´ Lada, Knı´zˇecı´ Plańeˇ, avenue of old trees along

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