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The saxicolous, xanthone-containing species of the genus s.l. (Physciaceae, ) in the Iberian Peninsula

Article in Nova Hedwigia · November 2009 DOI: 10.1127/0029-5035/2009/0089-0321

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The saxicolous, xanthone-containing species of the genus Buellia s.l. (Physciaceae, Ascomycota) in the Iberian Peninsula

by

Mireia Giralt1, Graciela Paz-Bermúdez2 and John A. Elix3 1Departament de Bioquímica i Biotecnologia (Àrea de Botànica), Facultat d’Enologia de Tarragona, Universitat Rovira i Virgili, Marcel·lí Domingo s/n, 43007, Tarragona, Spain e-mail: [email protected] 2E.U.E.T. Forestal, Campus A. Xunqueira, Universidade de Vigo, 36005, Pontevedra, Spain e-mail: [email protected] 3 Research School of Chemistry, Building 33, Australian National University, Canberra, ACT 0200, Australia. e-mail: [email protected]

With 3 figures

Giralt, M., G. Paz-Bermúdez & J.A. Elix (2009): The saxicolous, xanthone-containing species of the

genus Buellia s. l. (Physciaceae, Ascomycota)eschweizerbartxxx in ingenta the Iberian Peninsula. - Nova Hedwigia 89: 321– 334. Abstract: Five saxicolous species of Buellia s.l. containing xanthones are reported from the Iberian Peninsula: B. caldesiana, B. concinna, B. indissimilis, B. ocellata and a new species described here as B. vandenboomii. The latter also occurs in the Canary Islands (Tenerife). The status of B. indissimilis is clarified and an additional locality is reported for this rare species. The new combination Tetramelas concinnus is proposed. Key words: Buellia s. l., Tetramelas, Physciaceae, , , chemistry.

Introduction

The species treated in this study are characterized by containing xanthones. These lichen substances are concentrated in the thallus, the proper exciple and the epihy- menium. As a consequence the thallus frequently has a yellowish coloration, usually reacts C+ and KC+ orange and habitually fluoresces yellow, orange or pinkish under long wave (c. 350 nm) ultraviolet light, and the disc of the apothecia appears more or less pruinose (especially when young and wet). The thallus, proper exciple and epihymenium are often interspersed with golden yellow crystals of irregular shape and size. These crystals are clearly visible under polarized light and dissolve after treatment with C giving an orange reaction (microscope slide!).

DOI: 10.1127/0029-5035/2009/0089-0321 0029-5035/09/0089-0321 $ 3.50 © 2009 J. Cramer in der Gebrüder Borntraeger Verlagsbuchhandlung, D-14129 Berlin · D-70176 Stuttgart 321 All species treated are well distinguished from one another by several features and some are clearly related to species which lack xanthones: Tetramelas concinnus (Th. Fr.) Giralt (= B. concinna Th. Fr.) to the corticolous B. chloroleuca Körb., recently transferred into the genus Tetramelas (Nordin 2004); B. indissimilis to B. spuria (Schaer.) Anzi; and B. ocellata to B. aethalea (Ach.) Th. Fr.. There are no species which are obviously closely related to B. caldesiana and B. vandenboomii, although B. caldesiana appears similar in habit to B. tesserata Körb. According to Marbach (2000: 213), the content on arthothelin and the ascospore ornamentation relates B. caldesiana to the genus Gassicurtia Fée. The studied species are poorly known in the Iberian Peninsula and in the Mediterranean region and only occasionally reported from these areas. However, as the saxicolous European species of Buellia containing xanthones were studied previously by Scheidegger & Ruef (1988) and Scheidegger (1988, 1993), we only mention the main diagnostic and distinguishing characters of those species. Other known saxicolous, xanthone-containing Buellia s. lat. taxa include B. manami- ana Diederich (Aptroot et al. 1997), B. aeruginosa Nordin, Owe-Larss & Elix (Nordin et al. 1999), B. halonia (Ach.) Tuck., B. mamillana (Tuck.) W.A.Weber, B. sub- aethalea de Lesd., B. trachyspora Vain. (Bungartz et al. 2004a), B. vernicoma (Tuck.) Tuck. (Nordin 2000), B. georgei Trinkaus, H.Mayrhofer & Elix (Trinkaus et al. 2001), B. molonglo U.Grube & Elix (= B. halonia s. lat., Bungartz com. pers.) and B. cinnabarina U.Grube (Grube et al. 2004). According to Bungartz et al. (2004b), B. prospersa (Nyl.) Riddle (= Amandinea lecideina (H.Mayrhofer & Poelt) Scheid. & H.Mayr-hofer) may sometimes contain 4,5-dichloro-3-O-methylnorlichexanthone, thuringione, 4,5-dichlorolichexanthone and 2,4,5-trichlorolichexanthone while eschweizerbartxxx ingenta B. sequax (Nyl.) Zahlbr. may contain arthothelin. Although both species are present in the study area, they are not included in this contribution because we were unable to confirm these observations. More abundant are the known corticolous/lignicolous Buellia s. lat. species containing xanthones (See Kalb & Elix 1998; Marbach 2000; Nordin 2000). For terricolous/muscicolous species see Trinkaus et al. (2001).

Material and methods

The study is based on herbarium material from BCN, FI, GZU, H, LEB, MUB, PO, SANT, VAB, W, and from the private herbaria of M.Brand (Belgium) and P.v.d.Boom (The Netherlands). Material of B. concinna Th. Fr., B. jugorum (Arnold) Arnold and B. ocellata (Flot.) Körb. housed in GZU has been used for comparison. Specimens were examined by standard techniques using stereoscopic and compound microscopes. Current mycological terminology generally follows Kirk et al. (2001). Only free ascospores and conidia have been measured. Measurements were made in water at 1000× magnification. For the new species B. vandenboomii, mean value (M) and standard deviation (SD) were calculated and the results are given as (minimum value observed) M ± SD (maximum value observed). M, SD and n (the total number of ascospores measured) are given within parentheses. The terminology used for the asci follows Rambold et al. (1994), for the proper exciple and ascospore- wall ornamentation (Bungartz et al. 2004a), while for the ascospore-type and ontogeny that of Giralt (2001) and Bungartz et al. (2007) is followed.

322 Chemical constituents were identified by standardized thin-layer chromatography (TLC) (e.g. Culberson & Ammann 1979, Culberson et al. 1981, Culberson & Johnson 1982) and by high performance liquid chromatography (HPLC) (Elix et al. 2003).

Key to species

1 Thallus K+ red: norstictic acid present; UV+ bright orange...... B. indissimilis 1* Thallus K-: norstictic acid absent; UV+ orange...... 2 2 Apothecia lacking green or olivaceous, N+ red-violet pigment; young ascospores with callisporoid thickenings; ascospores occasionally becoming triseptate with age; paraphyses with strongly enlarged apical cells [to 7 μm wide]...... Tetramelas concinnus 2* Apothecia with green to olivaceous, N+ red-violet pigment in the epihymenium, hypothecium or proper exciple; young ascospores without callisporoid thickenings, never becoming triseptate with age; paraphyses with weakly enlarged apical cells [to 5 μm wide]...... 3 3 Ascospores smooth, (9–)10.5–12.5(–14) × (6–)6.5–7.5(–8.5) μm...... B. vandenboomii 3* Ascospores ornamented, significantly larger...... 4 4 Medulla I+ violet; ascospores (12–)13–16.5 × (7–)7.5–8(–9) μm, at least when young Physconia- type, not constricted at septum, microfoveate...... B. caldesiana 4* Medulla I-; ascospores (12–)13–16(–19) × (6–)7–8(–10) μm, always Buellia-type, constricted at septum, rugulate...... B. ocellata

Results and discussion

Buellia caldesiana Bagl. Fig. 1A Comment. Soc. Crit. Ital. 1(1): 19 (1861).

eschweizerbartxxx ingenta TYPI: ITALY, Liguria, nella valletta di S. Tecla, presso Genova, Baglietto, in Erbar. Crittogam. Ital. N° 88(1088) (FI- 5039(3) - LECTOTYPUS!: designated here; MOD - ISOLECTOYUS; TO - ISOLECTOYUS; W - ISOLECTOYUS!). ITALY, Liguria, nella valleta di Granarolo, Genova, Baglietto (MOD - SYNTYPUS).

NOMENCLATURAL NOTE: Scheidegger & Ruef (1988) reported that the isotypes of B. caldesiana were the specimens in TO and W, but this is not an effective typification under Art. 9.8 of the ICBN (McNeill et al. 2006) because an isotype presupposes a holotype and this one was not selected by Baglietto (1961). Further, as Baglietto (op. cit.) mentions two localities in his original diagnosis (‘valleta di S. Tecla’ and ‘valleta di Granarolo’, both near Genova), the specimens from la ‘valleta di Granarolo’ are syntypes. It should be noted that the isolectotype of B. caldesiana kept in TO has not been lend and that the results concluded after the material from MOD are based on photographs of the specimens and their labels. All the specimens collected in S. Tecla were distributed by Baglietto under number 88(1088) of the exsiccata Erbaria Crittogamico Italiano. It is likely the presence of more isolectotypes in other herbaria having been distributed under this exsiccata. A single syntype from Granarolo was located in MOD. Thallus crustose to slightly placodioid and subeffigurate at margin, thick, rimose to rimose-areolate, pale grey or cream, somewhat pruinose. Areolae with irregular surface. Prothallus absent. Medulla amyloid, I+ violet. Apothecia cryptolecanorine to subimmersed, usually surrounded by a thin, cracked thalline collar, rarely lecideine, up to 0.7 mm diam. Proper margin thin or absent. Disc plane to subconvex, usually covered with rough, greyish pruina. Proper exciple aethalea-type, poorly developed, totally interspersed with crystals C+ orange, colourless, nearly indistinct laterally,

323 Fig.1. Ascospore ontogeny and variability of:eschweizerbartxxx A. ingenta Buellia caldesiana, (BCN); B. Tetramelas concinnus (Hafellner-10988, GZU). Scale =10 μm. expanded to 20–30(–50) μm above and pigmented like the epihymenium. Hymenium colourless, without oil droplets, sometimes pale greyish in the lower part, 60–80 μm high. Epihymenium brown, olivaceous to dark grey-olivaceous, N+ red-violet, with abundant crystals, C+ orange. Hypothecium dark reddish brown, paler in the upper part. Apical cells of the paraphyses 3–4 μm diam., brown pigmented. Ascospores Physconia-type when young, with slight inner septal wall thickenings, becoming Buellia-type when mature (= +/- Physconia-type), (12–)13–16.5 × (7–)7.5–8(–9) μm, not constricted at septum, torus absent, ontogeny of type A, microfoveate (Fig. 1A). Conidia not seen.

CHEMISTRY: Thallus K+ yellow to yellow-orange, C+ pale orange, KC+ deep orange, PD-, UV+ pale orange-pink. Contains arthothelin [major], atranorin [minor], thiophanic acid [trace] and 4,5-dichloronorlichexanthone [trace].

HABITAT AND DISTRIBUTION: In the study area B. caldesiana occurs at 250–300 m altitude in the Prelitoral Mountains of Eastern Catalonia and the Valencian Country. It grows on exposed surfaces of meteorized granite. The Catalan record was already mentioned in Barbero (1998, ined.). It constituted the fourth record for B. caldesiana

324 which before that it was only known with certainty from the two Mediterranean type localities in northern Italy (Genova) designated by Baglietto (1861), and from one locality in Southern France (Hérault, Montpellier, PC-Weddel) mentioned by Scheidegger & Ruef (1988). There is an additional record from Sardinia (Baglietto 1879, sub. var. subpruinosa Bagl.) which hitherto has not been verified (Nimis 1993) and two additional specimens in MOD from “Bitiers” (France?) and Sarracco (Liguria?) which couldn’t be checked. The records mentioned below from the Valencian Country (VAB) and from Italy (FI) are new for the species.

OBSERVATIONS: Buellia caldesiana is characterized by its thick, pale greyish or creamy, somewhat chalky thallus which becomes slightly placodioid and subeffigurate along the margins; the presence of atranorin and arthothelin; the amyloid medulla (I+ violet); the persistent cryptolecanorine apothecia usually surrounded by a cracked thalline collar; the roughly pruinose discs; the green-pigmented, N+ red- violet epihymenium and the Buellia-type ascospores, showing Physconia-type thickenings when young (= +/- Physconia-type) and with microfoveate ornamentation. The microfoveate ornamentation can readily be recognized in overmature ascospores where tiny depressions can be observed instead of the prominences that are so typical for the rugulate ascospores. In the study area B. caldesiana could only be mistaken for B. tesserata, because of their somewhat similar habit. Nevertheless, the latter species lacks xanthones and shows a clearly effigurate to sublobate and fimbriate thallus margin (see Rico et al. 2003). In fact, one of the specimens studied was identified by Scheidegger as B. tesserata (under B. fimbriata).

ADDITIONAL SPECIMENS EXAMINED: ITALY, Liguria, Genova, 6.1855, Erbario O.Beccari (FI-5039(1)). Liguria, pr. Genova, Cengia Sambo, 10-XI-1947eschweizerbartxxx (FI-5039(2)). ingenta SPAIN, Catalunya, Barcelona, Maresme, prop de Dosrius, Can Llop (davant de la fábrica “Euroquímica”), c. 300 m, 28.I.1985, Barbero 17, 17.1 (BCN). País Valencià, Castelló, Villafamés, San Miquel, 500 m, 6.IX.1993, Calatayud (VAB- 7334). València, Puçol, El Picaio, 350 m, 26.II.1992, Calatayud (VAB-2627). València, Torres Torres, 250 m, 17.IV.1993, Calatayud (VAB-7115).

Buellia indissimilis (Nyl.) de Lesd. Fig. 2A Lich. Mexique: 26 (1914).

TYPUS: PORTUGAL, Porto, Lavadores, 1880, Newton (H-Nyl. 9310 - LECTOTYPUS!; H-Nyl. 10658 - ISOLECTOTYPUS!; PO-2832L - ISOLECTOTYPUS!).

NOMENCLATURAL NOTE: The specimen H-Nyl. 9310 was designated as the lectotype of Buellia indissimilis by Scheidegger & Ruef (1988). The same authors and Scheidegger (1993) report the specimen H-Nyl. 10568 as an isotype of B. indissimilis. According to the Art. 9.8 of the ICBN (McNeill et al. 2006), when a lectotype has been designated there are no isotypes but isolectotypes. Thus, the specimen H-Nyl. 10658 is an isolectotype. Thallus crustose, thin, rimose-areolate to areolate, whitish to cream. Medulla amyloid, I+ violet. Hypothallus usually distinct between areolae and surrounding the thallus, black, N+ red- violet. Apothecia cryptolecanorine, surrounded by a rather persistent thalline veil, becoming lecideine, sessile, broadly attached, 0.4–0.5(–0.7) mm diam. Proper margin at first hidden by the thalline veil, later clearly visible, narrow and slightly prominent. Disc plane, black, epruinose to moderately pruinose (better seen when wet). Proper exciple aethalea-type, nearly indistinct laterally, expanded to

325 Fig. 2. Ascospore ontogeny and variability of: A. Buellia indissimilis (PO-ISOLECTOTYPUS); B. B. ocellata (PO-2356L); C. B. vandenboomii (HOLOTYPUS). Scale = 10 μm.

50–60 μm above, usually brown pigmented,eschweizerbartxxx ingenta rarely greenish N+ red-violet, totally interspersed with crystals. Hymenium hyaline, without oil droplets, 50–65 μm high. Epihymenium brown, without or only with a few crystals. Hypothecium reddish brown, paler or hyaline in the upper part, up to 150 μm deep. Paraphyses with apical cells (2–)3–5 μm diam., brown pigmented. Asci Bacidia-type. Ascospores Physconia- type when young, with ephemeral, slight inner septal wall thickenings, Buellia-type when mature (= +/- Physconia-type), (9–)11–12(–13) × (5–)6–7 μm, not constricted at septum, torus absent, ontogeny of type A, smooth (Fig. 2A). Conidia bacilliform, (5–)6–7 × 1–1.2 μm.

CHEMISTRY: Thallus K+ red, C-, KC-, PD+ orange, UV+ bright orange. Contains norstictic acid [major], connorstictic acid [minor], 4,5-dichlorolichexanthone [minor], 5-chlorolichexanthone [minor or trace], 4-chlorolichexanthone [minor or trace], 4,5-dichloronorlichexanthone [trace] and 4,5-dichloro-3-O-methylnorlichexanthone [trace].

HABITAT AND DISTRIBUTION: Previously the silicicolous B. indissimilis was known only from the type locality on the coast of northern Portugal. The new record reported here is from the same area but 25 km further north, both localities in the Eurosiberian Region of the Iberian Peninsula. Sampaio (1921) reports a further locality but this collection could not be located.

OBSERVATIONS: Buellia indissimilis is characterized by the presence of norstictic acid in addition to several xanthones [not detected by C spot tests but readily observed

326 under UV light], the amyloid medulla, the flat apothecia usually surrounded by a white, thalline veil and the small, smooth, Buellia-type ascospores, of 9–12 × 5–7 μm, with slight septal wall thickenings corresponding to the Physconia-type when young (= +/- Physconia-type). Its conidia are here reported for the first time. It should be noted that while the width of the spores for this species is correctly reported in Scheidegger & Ruef (1988) [6.1–6.9 μm], it is erroneous in Scheidegger (1993) [6–9 μm]. Buellia indissimilis is closely related to B. spuria and in the study area it can only be mistaken for that species. However, the chemistry and the spore size and type readily distinguish both taxa. Buellia spuria lacks xanthones and has larger ascospores up to 15(–18) μm long, without Physconia inner wall thickenings. In addition, its epihymenium is green to olivaceous (N+ red-violet) and the hypothecium is dark brown. Among all saxicolous, xanthone-containing Buellia s.lat. species hitherto described (see the present introduction), B. indissimilis can only be mistaken for B. mamillana, a saxicolous species with a subtropical to tropical distribution containing also xanthones and norstictic acid. Nevertheless, B. mamillana has a different apothecial ontogeny and anatomy (initially lecanorine), norstictic acid does not constitute the major secondary lichen substance, its ascospores are larger and rugulate and its conidia are filiform, of 6–14 × 1–1.5 μm. (compare Bungartz et al. 2004a; 2007). Another saxicolous Buellia species which contains xanthones together with norstictic acid is B. prospersa but, in contrast to B. indissimilis, B. prospersa contains norstictic acid only in traces, its ascospores are Physconia-type at all ontogenic stages and its conidia are filiform (> 15 μm long). eschweizerbartxxx ingenta

ADDITIONAL SPECIMEN EXAMINED:PORTUGAL, Douro Litoral, Póvoa do Varzim, entre Amorim e Abremár, nos penedos, 1920, Sampaio 2227 (PO).

Buellia ocellata (Flot.) Körb. Fig. 2B Syst. Lich. Germ.: 224 (1855).

TYPUS: GERMANY, Baden Württemberg, Steinbül, Kemmler (BM - NEOTYPE). Thallus thin, areolate, lemon-yellow, yellowish-green to pale grey. Areolae contiguous or rarely scattered, small, flat, smooth to wrinkled. Hypothallus usually distinct between areolae and surrounding the thallus, black, N+ red-violet. Medulla non- amyloid, I-. Apothecia persistently cryptolecanorine, rounded to angular, up to 0.4 mm diam. Proper exciple aethalea-type, nearly indistinct and colourless laterally, expanded to 10–30 μm above and brown to green coloured (N+ red-violet), inter- spersed with a few crystals. Disc plane, black, epruinose. Hymenium colourless, without oil droplets, 90–100 μm high. Epihymenium brown or olivaceous to greenish (N+ red-violet), without crystals. Hypothecium brown, upper part grey olivaceous to greenish and N+ red-violet. Paraphyses with apical cells weakly enlarged to 3(–4) μm diam. and slightly pigmented. Asci Bacidia-type. Ascospores Buellia- type, (12–)13–16(–19) × (6–)7–8(–10) μm, constricted at septum, without torus, ontogeny of type A, rugulate. Conidia bacilliform, up to 7 μm long.

327 CHEMISTRY: K+ yellow to yellow-orange, C+ orange, KC+ deep orange, PD-, UV+ bright orange (TLC not done). Arthothelin ± stictic acid (Foucard et al. 2002). None of the specimens examined reacted PD+ orange or formed stictic acid crystals after the application of this reagent under the microscope. Thus, they most probably belong to the population lacking this secondary lichen substance.

HABITAT AND DISTRIBUTION: In the study area Buellia ocellata was hitherto only known in the Iberian Peninsula from two Atlantic localities, one in Galicia (NW Spain) and one in Portugal. It grows on siliceous rocks (including mafic rocks) from sea level to 500 m, in the coline belt. The material reported as B. ocellata by Scheidegger & Ruef (1988) from the central mountains of the Iberian Peninsula, proved to be B. aethalea when re-examined. The record from La Gomera (Canary Islands) in Scheidegger & Ruef (1988) has not been verified.

OBSERVATIONS: Buellia ocellata is a very uniform species, characterized by the thallus usually well delimited by a dark hypothalline line, the non-amyloid medulla, the small, persistently cryptolecanorine apothecia with an indistinct proper exciple and greenish upper part of the hypothecium (N+ red-violet), and its rather large Buellia- type ascospores which are constricted at the septum and have a rugulate ornamentation. It seems to be closely related to the xanthone-deficient B. aethalea. Studies using DNA sequences and phenotypic data (Helms et al. 2003, Nordin & Tibell 2005) support the close phylogenetic relationship between these species. Buellia jugorum (Arnold) Arnold, another xanthone-containing species not present in the study area, is distinguished from B. ocellata by the better developed thallus, the larger conidia and the alpine distribution (Foucard et al. 2002).

SPECIMENS EXAMINED: PORTUGAL, Minho, Póvoa de Lanhoso, entre Barreiros e Rendufinho, 1919, Sampaio 2368 (PO). Minho, Póvoa do Lanhoso,eschweizerbartxxx ingenta entre S. Gens e S. Roque, 1919, Sampaio 1950 (PO). Douro Litoral, Vila do Conde, nos penedos, 1920, Sampaio 2356 (PO). SPAIN, Galicia, A Coruña, Capelada, Cedeira, Coto de Outerio, 454 m, 15.IV.1995, Sánchez-Biezma & López de Silanes 9008 (SANT).

Buellia vandenboomii Giralt & Brand sp. nov. Figs 2C & 3 Mycobank no. MB514064 Thallus saxicola, autotrophicus vel parasiticus in thallis lichenum, ad 0.3 mm crassus, crustaceus, areolatus, flavescens. Medulla intense I+ caerulescens. Apothecia erumpentia vel sessilia, lecideina, marginibus propriis tenuibus, excipulum typo ‘aethalea’, pigmentum aeruginosum continens, ascosporae uniseptatae (9–)10.5–12.5(–14) × (6–)6.5–7.5(–8.5) μm [M = 11.5 × 7 μm], non ornatae, typo Physconia. Conidia bacilliformia, 4–5 × 1 μm. Thallus isoarthothelinum [major], 2,5-dichloro- norlichexanthonum [minor], acidum thiophanicum [paulum], 2,7-dichloronor-lichexanthonum [paulum], continens.

TYPUS: MACARONESIA, Canary Islands, Tenerife, Montañas del Teno, near col in the Santiago-Masca road, 16°49.4'W, 28°13.5'N, on Miocene volcanic rocks in open Euphorbia-Chamaecytisus vegetation, 900 m, 14.IV.1986, A.M.Brand 13681 (GZU- HOLOTYPUS) Thallus saxicolous, autotrophic or parasitic on other crustose , c. 0.3 mm thick, areolate. Areolae up to 2 mm diam., yellow, flat to subconvex, smooth, matt, loosely attached, margin dark green (N+ red-violet), contiguous, forming small rosette-like thalli up to 1 cm diam., or scattered. Cortex 30–60 μm thick, algal layer 40–60 μm thick, medulla amyloid, I+ violet, up to 200 μm thick. Apothecia initially

328 Fig. 3: Habit of Buellia vandenboomii (HOLOTYPUS) showing the thallus composed of thick, loosely attached areolae with dark green margins and the sessile, lecideine apothecia, which are initially erumpent and have thalline material attached to the margin. Scale = 0.75 mm.

eschweizerbartxxx ingenta erumpent with thalline material attached to the margin, then becoming adnate to sessile, lecideine, (0.2–)0.3–0.4(–0.6) mm diam., 1–3(–4) per areolae. Proper margin thin, black, excluded in convex apothecia. Disc flat to convex, covered by a white to yellowish pruina at least in young apothecia. Proper exciple aethalea-type, of 20–40(–60) μm thick, totally interspersed with C+ orange crystals; inner excipular hyphae narrow, colourless to pale brown (textura oblita), outer excipular hyphae of short, +/-swollen, brownish green cells, N+ red-violet. (textura oblita). Hymenium colourless, greenish in the upper part, without oil droplets, 60–80(–90) μm high. Subhymenium olivaceous grey to greenish. Epihymenium olivaceous grey to greenish, N+ red-violet, with abundant crystals giving a C+ orange reaction. Hypothecium dark reddish brown, paler in the upper part, 100–190 μm deep. Paraphyses with apical cells 3–4(–5) μm wide and pigmented dark brown. Asci Bacidia-type, 8-spored. Ascospores Physconia-type with inner septal wall thickenings, of (9–)10.5– 12.5(–14) × (6–)6.5–7.5(–8.5) μm (M = 11.5 × 7; SD = 1; 0.5; n = 72) not constricted at septum, without torus, very dark brown pigmented, ontogeny of type A, smooth at 1000 × (Fig. 2C). Conidia bacilliform, 4–5 × 1 μm. CHEMISTRY: Thallus K+ yellow-orange, C+ orange and KC+ deep orange, PD-, UV+ orange. Contains isoarthothelin [major], 2,5-dichloronorlichexanthone [minor], thiophanic acid [trace] and 2,7-dichloronorlichexanthone [trace].

329 ETYMOLOGY: The species is named in honour of Dr. Pieter G. van den Boom (Son, Netherlands), who has provided us with important collections and made significant contributions to lichenology.

HABITAT AND DISTRIBUTION: Buellia vandenboomii is only known from four localities. In the Iberian locality in the coastal mountains of southeastern Spain (Cabo de Gata), this species occurs in the Thermomediterranean belt of the Mediterranean region with semiarid ombroclimate. It was growing on Dimelaena radiata (Tuck.) Müll. Arg. and directly on volcanic acid rocks. Although a lot of material of D. radiata from the same and closed areas was checked to carry out the studies presented in Rico et al. (2003; 2006), only the specimen mentioned below was detected. In the Macaronesian localities, all situated in the mountains, the species occurs from middle to rather high altitude (570–900 m) growing associated with several lichens species or more often directly on volcanic rock.

OBSERVATIONS: The facultatively lichenicolous, well developed, thick, areolate, yellow thallus makes Buellia vandenboomii a quite conspicuous lichen species. Further, it is characterized by an amyloid medulla, lecideine apothecia, initially erumpent and pruinose, proper exciple, epihymenium and upper part of the hymenium tinged with the aeruginose pigment reacting N+ red-violet, and the smooth Physconia-type ascospores, very dark brown pigmented when mature. Chemically, the species is close to B. halonia, a widely distributed species in the Pacific coast of North America and the Southern Hemisphere, with larger ascospores of 11.5–19 × 6–9 μm (Bungartz et al. 2004b). Among the species treated B. vandenboomii can only be mistaken for B. caldesiana. Nevertheless, the latter species possesses significantly larger ascospores with a microfoveate ornamentation.

eschweizerbartxxx ingenta Other Buellia species facultatively lichenicolous on a wide range of hosts lichens, or lichenized and directly growing on rock, are B. badia (Fr.) A.Massal., T. concinnus and B. jugorum (Hafellner 2004). Buellia badia is clearly distinguished from B. vandenboomii by its brown thallus without secondary lichen substances. For differences between the two latter species and B. vandenboomii see under Tetramelas concinnus and B. ocellata.

ADDITIONAL SPECIMENS EXAMINED:MACARONESIA, Canary Islands, Gran Canaria, S of Las Palmas, S of Tafira Alta, small mountain ‘Bandama’, mixed trees, shrubs and outcrops on slopes along road, 15° 27.50'W/28° 02.30'N, 570 m, 6.II.2005, van den Boom 34157 (Hb.v.d.Boom). Tenerife, Montañas del Teno, near col in road Santiago-Masca, 900 m, 14.IV.1986, Brand 13682, 13683 (Hb. Brand - topotypes). Tenerife (NW), S of Santiago del Teide, NW of Tamaimo, path NE side of Montaña Guama, low exposed outcrops and vertical steep NW exposed outcrops in small valley, 16°49.14' W, 28°16.26'N, 630 m, 9.V.2007, van den Boom 37717 (Hb.v.d.Boom). SPAIN, Andalusia, Almeria, Sierra del Cabo de Gata, Las Madriguericas (Carboneras), WF9494, 180 m, 1.I.1973, Llimona (BCN-Lich).

Tetramelas concinnus (Th. Fr.) Giralt comb. nov. Fig. 1B Mycobank no. MB 514065

BAS.: Buellia concinna Th. Fr., Nova Acta Reg. Soc. Scient. Upsal., ser. 3(3): 232 (1860).

TYPUS: NORWAY, Finmark, Varanger, Nesseby, 1857, Th.M.Fries (UPS-LECTOTYPUS; GZU- ISOLECTOTYPUS!).

330 Thallus areolate, cream, yellowish or greenish yellow. Areolae dispersed to contiguous, flat to granulose-bullate, medulla amyloid or not, I+ strong violet, rarely I-. Prothallus absent. Apothecia lecideine, subimmersed to soon sessile and prominent, up to 0.75 mm diam. Proper margin distinct, only excluded in very convex apothecia. Disc plane to strongly convex, black, rarely faintly covered with greyish to yellowish pruina. Proper exciple leptocline-type, well developed, uniformly brown throughout, (30–)50–70 μm, inner part interspersed with crystals. Hymenium colourless, without oil droplets but sometimes with rows of C+ orange crystals, 60–80 μm high. Epihymenium brown, with C+ orange crystals, particularly in young apothecia. Hypothecium dark reddish brown. Paraphyses with strongly swollen apical cells [up to 7 μm diam.], with brown apical caps. Asci of Bacidia-type. Ascospores Physconia- type, when young with tendencies to the Callispora-type, 13–21 × 6.5–9(–10) μm, often curved, constricted at septum and sometimes becoming 2–3-septate when mature, ontogeny of type A, microrugulate to rugulate (Fig. 1B). According to Bungartz et al. (2004a) the measurements of the spore walls under transmission electron microscopy (TEM) are the following: proper wall 0.37–0.5 μm thick; intermediate layer < 0.03 μm; and perispore 0.11–0.17 μm. Conidia bacilliform, 2–5 μm long.

CHEMISTRY: Thallus K+ yellow-orange, C+ orange, KC+ deep orange, PD-, UV+ orange. Contains arthothelin [major], thiophanic acid [trace], 4,5-dichloronor- lichexanthone [minor]. Several additional xanthones were detected by HPLC by Bungartz et al. (2004a).

HABITAT AND DISTRIBUTION: This species is widely distributed in the Northern Hemisphere, but hitherto it was only known from two localities in the Iberian Peninsula (Scheidegger & Ruef 1988; Calatayud 1998, ined). It grows on siliceous rocks in submontane to montane areas from the northeasterneschweizerbartxxx ingenta to the southeastern Mediterranean mountains as well as in the mountains of central Spain. The specimens of T. concinnus from South Spain (MUB-869, 6895, 6983), grow associated with Dimelaena oreina (Ach.) Norman. Probably for this reason they were erroneously identified by Scheidegger (1993) as B. imshaugii Hafellner, a obligately lichenicolous species on D. oreina.

OBSERVATIONS: The genus Tetramelas Norman was resurrected by Marbach (2000). Kalb (2004) and Nordin (2004) emended the genus adding the presence of 6-O- methylarthothelin and the relative thickness of the parts of the spore wall (proper wall > 1/2 thickness of the perispore) as diagnostic characters, respectively. Since then, twelve species have been combined into the genus and a new species has been described, namely the lichenicolous T. phaeophysciae A.Nordin & Tibell (Nordin & Tibell 2005). Among these Tetramelas species, T. triphragmioides (Anzi) A.Nordin & Tibell contains arthothelin rather than 6-O-methylarthothelin, T. papillatus (Sommerf.) Kalb contains atranorin rather than xanthones and T. pulverulentus (Anzi) A.Nordin & Tibell and T. phaeophysciae lack secondary lichen substances altogether. As it has been shown that Tetramelas constitutes a well founded segregate of Buellia s. lat. (Helms et al. 2003, Nordin & Tibell 2005), the new combination Tetramelas concinnus is proposed. Like most species in the genus, T. concinnus contains several xanthones (see above and Bungartz et al. 2004a), has ascospores which are often

331 curved, have pointed apices, rugulate walls and may become triseptate at maturity and paraphyses with strongly swollen apical cells. In our opinion, a further important feature of the genus Tetramelas is the presence of callisporoid thickenings in intermediate ontogenic stages of the ascospores [e.g. T. chloroleucus (Körb.) A.Nordin and T. insignis (Nägeli ex Hepp) Kalb as shown in fig. 1 (Giralt et al. 2000) and T. regiomontanus Marbach in fig. 159 (Marbach 2000)]. According to Kalb (2004), in addition to the presence of xanthones, other diagnostic characters for Tetramelas included the typically muscicolous habitat, large ascospores and the arctic-antarctic (or sub-alpine) distribution. In contrast, T. concinnus is saxicolous and has rather small ascospores as do T. granulosus (Darb.) A.Nordin and T. chloroleucus, both combined into Tetramelas by Nordin (2004). Tetramelas concinnus is the first species in the genus which occurs in montane to submontane areas of the Mediterranean region and which has Physconia-type ascospores that nevertheless show at least some ontogenetic affinities with the Callispora-type. Among the species treated in this contribution, T. concinnus is the only one with a thick, uniformly brown proper exciple, commonly curved ascospores with Callispora- type tendencies which may become triseptate with age, paraphyses with strongly swollen apical cells and total absence of the aeruginose N+ red-violet pigment. Among the specimens studied we have detected two populations of T. concinnus: one with 1-septate ascospores, of 14–17 × 6.5–8 μm with rugulate spore-walls and the other with larger ascospores (up to 21 × 9(–10) μm) which often become triseptate and with less prominently ornamented (= microrugulate) spore-walls. However, the presence of an intermediate specimen (BCN-754) and the absence of other correlating

characters have convinced us to maintaineschweizerbartxxx ingenta the two types within one taxon. Furthermore, other authors quote a broad range for the size of the ascospores of this species (Foucard et al. 2002, Bungartz et al. 2004a). All the specimens studied had an amyloid medulla except for the isolectotype (GZU).

ADDITIONAL SPECIMENS EXAMINED: SPAIN, Andalusia, Almeria, Tabernas, Sierra Alhamilla, Colativí, 1300 m, 21.IX.1979, Egea & Llimona (MUB-6895, 6973). Almeria, Bacares, Sierra de los Filabres, Merendera, 1984, 9.VIII.1978, Egea (MUB-869). Castella-Lleó, Lleó, Mascariel, 1600 m, 23.VII.1989, Terrón (LEB-207). Catalunya, Barcelona, Montseny, Mirador prop de Can Penyacanyes, 1200 m, sobre esquistos, 21.VI.1981, Hladun 754 (BCN-Lich). Girona, Port de la Selva, Torrent Puig Vaquer, vora urbanització Beleser, Font de la Clareta, c. 100 m, 21.I.1984, Hladun (BCN-Lich). Madrid, Sierra de Guadarrama, auffahrt von Lozoya auf den Puerto de Navafría, 1300 m, silikatblöcke, 7.IX.1980, Hafellner 10988 (GZU). País Valencià, Castelló, Montán, Los Agualiches, sobre arenisca roja del Buntsandstein, 680 m, 28.II.1997, Calatayud 7804, 7788dup1 (VAB).

Acknowledgements

The authors are indebted to the keepers of BCN, FI, GZU, H, LEB, MUB, PO, SANT, VAB and W for the loan of material used in this study, to Anders Nordin and Frank Bungartz for valuable comments on the manuscript and to Harry Sipman for correcting the Latin diagnosis. We also wish to thank Mauro Tretiach and the herbarium of Modena (MOD) for supplying information on Buellia caldesiana and Pep Agulló for improving the figures. The first author thanks the ‘Comissionat per a la Recerca’ (Catalan Government) and the second the University of Vigo for financial support.

332 References

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Received 10 December 2008, accepted in revised form 30 April 2009.

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