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ASPICILIA CRESPIANA, A NEW LICHEN SPECIES FROM SOUTHERN EUROPE
Lichenologist 31(2): 129–139 (1999) Article No. lich.1998.0181 Available online at http://www.idealibrary.com on
ASPICILIA CRESPIANA, A NEW LICHEN SPECIES FROM SOUTHERN EUROPE Vı´ctor J. RICO*
Abstract: Aspicilia crespiana Rico is described as a new species from central Spain and Sardinia (Italy). In central Spain it is common and has a well-defined ecology. It is found at middle to high altitudes (from 400 to 1900 m), overgrowing mosses on exposed and horizontal to more or less vertical siliceous rocks. Closely related to the A. contorta (Hoffm.) Kremp. complex, A. crespiana differs in its muscicolous habit and thallus development, consisting of a network of lichenized squamules interlinked by fungal rhizomorphs. 1999 The British Lichen Society
Introduction In recent years a distinctive, muscicolous, squamulose to subfruticose Aspicilia s. lat., with strongly developed rhizomorphs, has been collected frequently in central Spain. Additional material from Sardinia (Italy) was sent to the author by Prof. P. L. Nimis (Trieste). The term ‘ rhizomorph ’ is used here in its broad sense, referring to linear hyphal aggregates with irregular structure, indeterminate growth, and with a generally exploratory function, as well as serving in attachment and propagation. Lichen rhizomorphs lack organized apices. Further information on the use of this term in lichenized fungi is given by Sanders (1994), and in non-lichenized fungi by Cairney et al. (1991). This taxon has been referred to as ‘ Aspicilia crespiana Rico ined. ’ in some publications (Nimis 1993; Sanders & Rico 1992). As this lichen apparently does not agree with any taxa in previously published papers on aspicilioid lichens it is described here as a new species. Materials and Methods Seventy-eight specimens of the new species, from seven provinces in Spain and one in Sardinia (Italy) were investigated (material deposited in MA, MAF and TSB). Freezing microtome sections (SLEE-MTC) and hand sections were mounted in water or in lactophenol-cotton-blue (Merck Rf. 13741). To study ascus structure, Lugol’s solution was added to sections and squash preparations after pre-treatment with KOH. A Nikon Optiphot-AFX II compound microscope was used for micrographs of sections and a Zeiss 47 50 52 dissecting microscope was used for habit photographs. Thirty-five selected specimens were analysed by TLC, following White & James (1985). Special care was taken with ascospore measurements and only well-developed and mature ascospores were analysed. These are more frequent in young and medium-sized apothecia of the studied material. The dimensions and length:breadth ratio (l:b) of ascospores are based on 93 *Departamento de Biologı´a Vegetal II, Facultad de Farmacia, Universidad Complutense, E–28040 Madrid, Spain. E-mail: [email protected] 0024–2829/99/020129+11 $30.00/0
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readings of several specimens of Aspicilia crespiana and on 40 readings of 4 specimens of A. contorta subsp. contorta and of A. contorta subsp. hoffmanniana (10 readings for each specimen cited below). Extreme values have been placed between brackets where they represented no more than 10% of the spores. For comparison of spore shape between taxa, I followed the terminology and corresponding l:b ratio defined by Bas (1969: 321–322) [globose (l:b of 1·0–1·05), subglobose (1·05–1·15), broadly ellipsoid (1·15–1·3), ellipsoid (1·3–1·6), elongate (1·6–2·0), cylindrical (2·0–3·0), bacilliform (>3·0)]. Besides the selected material of A. crespiana reported below, the following selected specimens were examined for comparison: Aspicilia contorta subsp. contorta: Spain: Burgos, Morentı´n, carretera de Castil de Peones a Villafranca-Montes de Oca, valle de Oca, monumento al General Mola, c. 950 m, guijarros calizos, 1996, G. Renobales (MAF). Madrid: Pelayos de la Presa, cruce del rı´o Alberche con la carretera comarcal 501, 550 m, 30TUK8969, granito con impregnaciones de polvo calizo, 1984, E. Barreno & V. J. Rico 32/1 (MAF); Perales de Tajun˜a, 650–780 m, 30TVK65, rocas calizas, 1980, E. Barreno & A. Merino (MAF). Navarra: Zudaire (Amezcoa Baja), 640 m, en la base de Quercus faginea, 1984, B. Aguirre (MAF). Aspicilia contorta subsp. hoffmanniana: Spain: Cáceres, Toril, Parque Natural de Monfragüe, Sierra del Serrejón, entre el Poblado del Embalse y La Herguijuela, Portilla del Tietar, 300 m, 30STK4716, cuarcitas, 1988, V. J. Rico 2088/2 (MAF). Madrid: Pelayos de la Presa, cruce del rı´o Alberche con la carratera comarcal 501, 550 m, 30TUK8969, granito, 1984, E. Barreno & V. J. Rico 55/2 (MAF); Hoyo de Manzanares, cerca de la urbanización La Berzosa, 980 m, 30TVK2196, granito, 1987, F. L. Redondo, Rico 1225/4 (MAF). Vizcaya: Ceánuri, Monte Larralde, 900 m, 30TWN2165, caliza compacta, 1985, M. Herrera & G. Renobales 1412 (BIO).
The Species Aspicilia crespiana Rico sp. nov. Aspicilia contortae affinis est, sed habito muscicola, thallo squamuloso vel subfruticuloso cum rhizomorphis, apotheciis distincte majoribus et sporis plerumque ellipsoidalibus differt. Thallus squamulosus vel subfruticulosus; crescit super Grimmia sp. in rupibus siliceus. Squamulae diametro usque ad 5 mm, imbricatae, dense congestae, apicem versus ascendentes vel lobatae, subtus ad rhizomorphis colligatae. Rhizomorphae bene evolutae, usque ad aliquot cm. longitudine, diametro usque ad 1·5 mm, irregulariter inaequaliter ramosae, algae destitutae; inter Grimmia sp. foliis et caulis et inter rupibus rimae penetrantes. Apothecia communis, primum cryptolecanorinia deinde lecideinia vel lecanorinia, immersa vel sessilia, usque ad 3·5 mm diametro; margine incrassato. Asci clavati, 2- vel 6-spori; tholis crassis, I. Ascosporae incolorae, simplices, plerumque ellipsoideae, (11–)15–32(–45)(8–)11–20(–31) ìm, cum tenuis pariete. Sine acido lichenoso. Typus: Spain, Madrid, San Martı´n de Valdeiglesias, road from Cadalso de los Vidrios to Pelayos de la Presa, km 3, Corcobada, 740 m, overgrowing Grimmia sp. on horizontal sun-exposed granitic rocks, 30TUK843668, 12 February 1988, V. J. Rico 1249/1 & M. A. Florido (MAF-Lich. 4221—holotypus; MA-Lichen 3274—isotypus).
(Figs 1–3) Thallus squamulose to subfruticose, consisting of commonly imbricate or dispersed lichenized squamules interconnected by abundant mycobiont rhizomorphs, forming large patches up to several decimetres diam.; squamules overgrowing saxicolous moss cushions (generally Grimmia species) and soil and rock in the vicinity, the rhizomorphs penetrating among these mosses and through the surrounding soil and rock. Squamules (Figs 1A, 2) up to 5 mm wide, highly variable. Mature squamules on mosses and soil, irregularly imbricate to contiguous, frequently fused to each other laterally, orbicular to elongate, generally ascending mainly at the tips, surface flat to concave or strongly convex, margins sinuate to lobulate, lobules rounded or commonly
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F. 1. Aspicilia crespiana. A, squamulose to subfruticose thallus growing on Grimmia sp. (holotype, MAF); B, lower side of thallus, showing rhizomorphs between mosses, rock fissures and soil; C, transverse section through mature lecanoroid apothecium in lactophenol-cotton-blue (holotype, MAF). Scales: A–B=2 mm; C=200 ìm.
verrucose to cylindrical and sometimes divided. Squamules of thallus margins on exposed, moss-free rocks, rare to frequent, sometimes areole-like, appressed, contiguous to scattered, orbicular to elongated,
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F. 2. Aspicilia crespiana (holotype, MAF). A, detail of thallus showing mature squamules overgrowing mosses; B, lateral view of squamules interlinked with rhizomorphs; C, detail of a subfruticose lichenized squamule (C1) with irregularly branched mycobiont rhizomorph (C2). Scale: 5 mm.
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margins mostly rounded, surface convex to commonly verrucose or bullate to ascending. Prothallic structures occasionally present on squamule margins, grey to white fan-like bands on moss leaves, dark strap-like bands on exposed rocks. Upper surface matt, white to grey or partly olivaceous to brownish, slightly roughened or smooth, frequently with white maculae consisting of accumulations of dead cortical cells; pycnidia or apothecial primordia also common as white to brown or black spots, without pseudocyphellae and pruina. Upper cortex 18–75(–90) ìm thick, paraplectenchymatous, of rounded cells with lumina 2·5–10(–12) ìm wide, brown-green pigmented at the surface; epinecral layer variable in thickness, consisting of dead mycobiont cell walls, without crystals or granules; tips of the ascending squamules also corticated below. Photobiont green, chlorococcoid, unicellular, cells 5–13 ìm diam.; arranged in a continuous layer or in a denticulate one when thallus squamules are thicker. Medulla loose to paraplectenchymatous, with pachydermatous hyphae, occasionally with grey granules, highly variable in thickness; strands of meso- to pachydermatous hyphae crossing this layer in a prosoplectenchymatous arrangement, extending from the squamules into the rhizomorphs. Lower surface of mature squamules white to ochraceous, without cortex or with a rudimentary one of 1, 2 or 3 rows of isodiametric cells, continuous with those of the rhizomorphs. Rhizomorphs (Figs 1B, 2) delicate, lacking algae, pale whitish or yellowish to pink and later darkly pigmented, terete to flattened, variable in length (up to several cm) and thickness (to 1·5 mm), branching extensively without any discernible pattern, interconnecting squamules terminally or laterally and giving rise to new ones. Rhizomorphs possess a central cord of densely packed meso- to pachy dermatous hyphae, in a prosoplectenchymatous arrangement. Surrounding the longitudinally arranged strands there is a rudimentary cortex of isodiametric cells (paraplectenchymatous) with or without pigmented walls. Distally from the squamules, the rhizomorphs eventually become fine, ecorticate and diffuse. Apothecia numerous, up to 3·5 mm wide, simple or composite, orbicular to compressed, immersed to elevated above the squamule or commonly sessile with constricted base, rarely slightly peltate, one to several per squamule, sometimes regenerating; cryptolecanorine or nearly urceolate at first, becoming lecanoroid (Fig. 1C) or lecideoid in larger fruits. Disc concave to flat, black. Margin distinct, later prominent, concolorous with thallus (lecanoroid) to brownish or black when thalline margin is reduced (lecideoid). Excipulum propium paraplectenchymatous in outer part, with brown-green to blackpigmented cell walls at the surface; highly variable in development and thickness, reduced in immersed apothecia to extended and rarely flabellate in large lecideoid fruits. Hypothecium colourless, composed of rounded cells, mostly reduced, with or without an algal layer underneath. Hymenium 105–160(–175) ìm high, conglutinated, colourless, without oil droplets; epithecium olivaceous to greenish, in young apothecia covered by dead cortical cells; subhymenium colourless. Paraphyses (Fig. 3B) branched and anastomosing, moniliform to submoniliform, with 1 to 9 rounded cells at the tips, up to 7 ìm thick at the olive-coloured apices, conglutinated. Asci (Fig. 3C) clavate, (75–)105–14020–40 ìm, 2- to 6-spored, mostly
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F. 3. Aspicilia crespiana. A, ascospores; A1 mature spore joined with the wall of a degenerating one; B, paraphyses; C, mature asci containing 2–6 spores; C1, ascus after treatment with I. Scale: 20 ìm.
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4-spored; apical apparatus thick, K/I; with a fuzzy, amyloid coat (Fig. 3C1). Ascospores (Fig. 3A) unicellular, colourless, thin-walled, rapidly degenerating at maturity, (11–)15–32(–45)(8–)11–20(–31) ìm, l:b ratio (1·03–)1·33–1·94(–2·42), l:b ratio mean value of 1·58, (globose) ellipsoid to elongate (cylindrical), mainly ellipsoid, rarely ovoid. Pycnidia common, more frequent at tips of squamules, immersed; ostiole greenish-brown to black; wall hyaline, rounded to convoluted; conidia 7–121 ìm, bacilliform, straight. Chemistry. No lichen substances detected by TLC; cortex and medulla K, C, KC, PD and I; pigment of epithecium and ostiolar area of pycnidia HNO3 + emerald green (Aspicilia-green). Etymology. This species is dedicated to Luis Crespı´ Jaume (Madrid, 1889–1963), who collected it during the nineteen twenties (cf. Crespı´ & González Bueno 1990). Distribution and habitat. Currently, A. crespiana is known from one locality in Sardinia (Italy) and from 14 localities in central Spain, where it is common and forms large, rather conspicuous patches. In A u vila (840 m alt.), Badajoz (725 m), Cáceres (400 m), Madrid (550–1300 m), Segovia (1130 m) and Toledo (580 m) provinces (Spain), it was collected in dry to subhumid situations from the mesomediterranean to supramediterranean belts of the Mediterranean Region (Rivas-Martı´nez 1987). It characteristically overgrows Grimmia species on siliceous rocks, mainly granite, usually in widely open situations amongst Quercus pyrenaica, Q. rotundifolia, Q. suber or cultivated Pinus pinea or P. sylvestris forests. In these localities a Cistus understorey is common. Aspicilia crespiana is found on sunny, horizontal to vertical rock surfaces, frequently with small amounts of sandy soil below the overgrown moss. The material from the Spanish province of Palencia (1900 m) is exceptional, having been collected in the subalpine belt of the Eurosiberian Region (Rivas-Martı´nez 1987), on exposed rock surfaces accompanied by few conspicuous gametophytes of an unidentified bryophyte. The most frequent lichens co-occurring with A. crespiana in central Spain are: Lecanora aff. muralis, Lepraria neglecta, Neofuscelia pulla, N. verruculifera, Xanthoparmelia tinctina and sometimes species of Cladonia and Lasallia pustulata. When growing near small rivers or raintracks, in slightly humid situations, Leptochidium albociliatum and Polychidium muscicola may also occur close to A. crespiana. Buellia badia and a taxon of the Rhizocarpon viriciatrum group are also common lichens growing on A. crespiana thalli. They together comprise a clearly differentiated community on mosses overgrowing horizontal or inclined, sun-exposed siliceous rocks. Remarks. The new species is easily recognized in the field by its primary muscicolous habit, and thallus development. Aspicilia crespiana is comprised of two interconnected structures, lichenized squamules and mycobiont rhizomorphs. The two types of structures can, under suitable conditions, give rise to each other. A detailed analysis of thallus development and variability, lichenizing rhizomorphs and substratum occupation of the new species is given by Sanders & Rico (1992). Similar types of structures have also been
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observed in other squamulose lichens, such as the terricolous Endocarpon pusillum (Malone 1977; Wagner & Letrouit-Galinou 1988) or the saxicolous Lecanora rhizinata (Barreno & Rico 1984; Sanders 1994). Terricolous species of Toninia frequently occur among mosses on soil, producing an extensive rhizomorphic system (Sanders 1994). The initial cryptolecanorine ascomatal structure, the ascus type with thick apical apparatus and amyloid gelatinous exoascal layer, moniliform paraphyses, olivaceous epithecium with HNO3 + reaction, and conidia morphology, confirms that this species belongs to the genus Aspicilia s. lat. (Clauzade & Roux 1984; Hafellner 1991; Purvis et al. 1992). The apothecial form, spore and conidial size, thickness of ascospore walls and number of spores per ascus, support its inclusion in Aspicilia subgen. Aspicilia (Clauzade & Roux 1984), and indicates that A. crespiana belongs to the A. calcarea (L.) Mudd group and is related to silicicolous forms of A. contorta (Hoffm.) Kremp. (Clauzade & Roux 1985; Esnault 1985; Ekman & Fröberg 1988; Hafellner 1991). Aspicilia contorta is a common lichen growing on calcareous rocks, limestone, mortar, siliceous rocks or rarely lignum of North Africa, Europe and North America (Clauzade & Roux 1985; Esnault 1985; Purvis et al. 1992). In Europe, it has a wide ecological amplitude and a correspondingly wide morphological variation (Nimis 1993). Several forms (Esnault 1985; Clauzade & Roux 1985) and intermediate specimens (Ekman & Fröberg 1988) have been described. For comparison, I follow Fröberg (1989) and Wirth (1995), who recognize two subspecies of A. contorta, A. contorta subsp. contorta and A. contorta subsp. hoffmanniana Ekman & Fröberg. Table 1 presents a comparison of the main taxonomic characters of A. crespiana and the two subspecies of A. contorta, based on recent, confirmed Spanish material (see Materials and Methods) and on bibliography. Aspicilia crespiana is clearly separated from A. contorta subsp. contorta and subsp. hoffmanniana by its muscicolous, squamulose to subfruticose thallus with strongly developed rhizomorphs. The apothecia and ascospore shape are also useful characters. Typically, the young apothecia of A. crespiana are similar to those (young and mature) of both subspecies of A. contorta, immersed, cryptolecanorine, with reduced excipulum propium and raised thallus margin. Mature apothecia of A. crespiana appear nearly urceolate, lecanoroid or lecideoid, and nearly all have a prominent margin. Moreover, a single mature apothecium of A. crespiana may have one part lecanoroid, with a prominent thalline margin and a reduced excipulum propium, and other parts lecideoid, with a prominent excipulum propium and a reduced or non-existent thalline margin. Sometimes, when apothecia are situated at the tips of ascending squamules or centred in convex ones, a short stalk can be distinguished. Despite the number of specimens examined, A. crespiana sporulated poorly, and only a limited number of mature and well-developed ascospores (93) could be measured. According to Bas (1969) the ascospores of A. crespiana are mainly ellipsoid in shape, and those of both subspecies of A. contorta are mainly broadly ellipsoid, as is inferred by comparing l:b ratio mean values (Table 1). Aspicilia contorta subsp. contorta and subsp. hoffmanniana occur together with the new species even in the same
0·5–2·5 mm diam., surface flat to convex, margins mainly rounded to lobulate
–
Up to 1·2 mm diam., immersed to subimmersed, cryptolecanorine to lecanoroid, margin later slightly raised, often pruinose
(20–)21–26(–30)(17–)18–22 (1·04–)1·09–1·33(–1·39) 1·20 (broadly ellipsoid)
Aspicilin ?*
Mainly on calcareous rocks, rarely on base enriched siliceous rocks or on lignum
N. C. & S. Europe‡
Areole/squamules
Rhizomorphs
Apothecia
Ascospore size (ìm) l:b ratio l:b ratio mean value
Chemistry
Habitat
Distribution (Europe)
N. C. & S. Europe‡
On calcareous or on siliceous rocks
Sometimes aspicilin ?*
(19–)20–30(–32)(15–)18–21(–25) (1·04–)1·1–1·55(–1·78) 1·28 (broadly ellipsoid)
Up to 1 mm diam., immersed to sessile, cryptolecanorine to lecanoroid, margin later raised to prominent, sometimes pruinose
–
0·1–1·8 mm diam., surface convex, margins mainly angled
Crustose, areolate, continuous to dispersed in thallus margins, grey to whitish, sometimes pruinose
subsp. hoffmanniana
C. Spain and Sardinia
Muscicolous and secondarily on siliceous rocks and soil in the vicinity
–
(11–)15–32(–45)(8–)11–20(–31) (1·03–)1·33–1·94(–2·42) 1·58(ellipsoid)
Up to 3·5 mm diam., immersed to sessile or slightly peltate, cryptolecanorine to lecanoroid or lecideoid, margin later prominent, never pruinose
+, Interconnecting squamules
Up to 5 mm diam., surface flat to concave or convex, margins with rounded, divided and cylindrical lobules, mainly ascending
Squamulose to subfruticose, continuous to dispersed in thallus margins, grey to whitish, never pruinose
A. crespiana
Aspicilia crespiana—Rico
*According to Ekman & Fröberg (1988). ‡According to Nimis (1993), Santesson (1993) and Wirth (1995).
Crustose, areolate to squamulose, dispersed in groups of scales, white to greyish, usually pruinose
subsp. contorta
Thallus
Characters
A. contorta
T 1. Main characters for distinguishing among Aspicilia contorta subsp. contorta, A. contorta subsp. hoffmanniana and A. crespiana
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locality, as seen in the studied material (Rico 32/1, Rico 55/2 and Rico 2968/1 respectively). However, it has a different habit in addition to the characters described above. Closely related to the A. contorta complex, and consequently with A. crespiana, are the fruticose, terricolous and vagrant species of the A. hispida Mereschk. agg. (Hafellner 1991). Morphologically, A. hispida is clearly different from the new species by its fruticose thallus, with cylindrical, pseudocyphellatae ramifications, lacking lichenized squamules and mycobiont rhizomorphs (Clauzade & Roux 1987). Additional selected specimens examined: Italy: Sardinia: Prov. Cagliari, Riserva Naturale di M. Arcosu, su muschi epilitici su trechite, 800 m, 1989, P. L. Nimis & M. Tretiach (TSB, MAF).—Spain: A u vila: Cebreros, en el km 1·4 de la carretera de Cebreros a El Hoyo de Pinares, ladera este del alto de El Castan˜ar, 840 m, 30TUK7680, sobre briófitos en granito, 1997, V. J. Rico 3329 (MAF). Badajoz: Garbayuela, Sierra de los Villares, a la derecha de la carretera desde Herrera del Duque, cerca de una cantera y antes del puerto de Los Carneros, 725 m, 30SUJ2829, sobre briófitos en cuarcitas y pizarras muy soleadas, 1996, V. J. Rico 3202 & M. A. Florido (MAF). Cáceres: junto a la Sierra de San Pedro, a 6 km por el camino que parte del suroeste de Malpartida de Cáceres, en las márgenes del rı´o Salor, 400 m, 29SQD1264, sobre briófitos en granito, 1992, F. J. Garcı´a González, Rico 2962 (MAF). Madrid: cerca de Hoyo de Manzanares, El Quemadillo, sobre musgos en pen˜as, 1926, L. Crespı´ (MA); San Martı´n de Valdeiglesias, embalse de San Juan: cerro Almodón, 650 m, 30TUK8273, sobre briófitos en granito, 1984, E. Barreno & V. J. Rico 114 (MAF); ibid., cerro Cabreruela, 690 m, 30TUK8573, sobre briófitos en granito, 1985, E. Barreno & V. J. Rico 334 (MAF); ibid., cerro de El Yelmo, 680 m, 30TUK8872, sobre briófitos en granito, 1991, V. J. Rico 2657 & M. A. Florido (MAF); Pelayos de la Presa, cruce del rı´o Alberche con la carretera comarcal 501, 550 m, 30TUK8969, granito, 1993, V. J. Rico 2968/1 (MAF); Cenicientos, pen˜a Cenicientos, 1253 m, 30TUK7259, 1986, J. M. Esteban, Manrique M34–40 (MAF); Robledo de Chavela, pico de El Almojón, 1175 m, 30TUK9581, sobre briófitos en granito, 1986, V. J. Rico 797 (MAF); San Lorenzo de El Escorial, Silla de Felipe II (bellow), on mosses over rock, exposed, 1991, W. B. Sanders (MAF); Lozoya, El Chaparral, 1300 m, 30TVL3634, sobre briófitos en gneis, 1986, V. J. Rico 744/1 & E. Manrique (MAF). Palencia: Resobada, pen˜a Curavacas, north face, 1900 m, 30TUN66, on siliceous conglomerate rock, no obvious bryophyte, 1992, W. B. Sanders (MAF). Segovia: San Ildefonso (La Granja), Parque de Santa Cecilia, cerca de la carretera de La Granja a Segovia, 1130 m, 30TVL1428, sobre briófitos en granito, 1998, V. J. Rico 3333/1 (MAF). Toledo: Nombela, en el km 13·3 de la carretera de Nombela a Pelahustán, 580 m, 30TUK6946, sobre briófitos en granito, 1997, V. J. Rico 3322 (MAF). I thank Dr C. Roux (Marseille) and Dr J. Esnault (Acigne) for their valuable comments on the specimens, Dr W. B. Sanders (San Francisco) for helping with the material and the photographs and for reviewing the manuscript, Dr G. Renobales (Vitoria) for the loan of Aspicilia contorta, Dr J. Pizarro (Madrid) for doing the line drawings, and Prof. P. L. Nimis (Trieste) for kindly sending me material from Sardinia. My dear friends Pepe Bella and Philip Mason are thanked also. R Barreno, E. & Rico, V. J. (1984) Sobre la biologı´a de los lı´quenes I. Anatomı´a, morfologı´a y estructuras vegetativas. Anales de Biologı´a (Murcia) 1: 161–195. Bas, C. (1969) Morphology and subdivision of Amanita and a monograph of its section Lepidella. Persoonia 5: 285–579. Cairney, J. W. G., Jennings, D. H. & Agerer, R. (1991) The nomenclature of fungal multi-hyphal linear aggregates. Cryptogamic Botany 2: 246–251. Clauzade, G. & Roux, C. (1984) Les genres Aspicilia Massal. et Bellemerea Hafellner et Roux. Bulletin de la Société Botanique du Centre-Ouest, Nouvelle série 15: 127–141. Clauzade, G. & Roux, C. (1985) Likenoj de Okcidenta Euˇropo. Ilustrita determinlibro. Bulletin de la Société Botanique du Centre-Ouest, Nouvelle série, numéro spécial 7-1985: 1–893.
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Clauzade, G. & Roux, C. (1987) Likenoj de Okcidenta Euˇropo. Suplemento 2a. Bulletin de la Société Botanique du Centre-Ouest, Nouvelle série 18: 177–214. Crespı´, A. L. & González Bueno, A. (1990) Botanicae historiae schedulae sparsae 5. Luis Crespı´ Jaume (Madrid, 1889–Madrid, 1963). Acta Botánica Malacitana 15: 343–345. Ekman, S. & Frö berg, L. (1988) Taxonomical problems in Aspicilia contorta and A. hoffmannii—An effect of hybridization? International Journal of Mycology and Lichenology 3: 215–225. Esnault, J. (1985) Le genre Aspicilia Mass. (Lichens) en Algérie: étude des caractères taxonomiques et de leur variabilité. Doctoral Thesis, Laboratoire de Botanique L. Daniel, Université de Rennes I. Fröberg, L. (1989) The Calcicolous Lichens on the Great Alvar of O } land, Sweden. Lund: Institutionen för Systematisk Botanik. Hafellner, J. (1991) Die Gattung Aspicilia, ihre Ableitungen nebst Bemerkungen über cryptolecanorine Ascocarpsorganisation bei anderen Genera der Lecanorales (Ascomycetes lichenisati). Acta Botánica Malacitana 16: 133–140. Malone, C. P. (1977) Observations on Endocarpon pusillum: the role of rhizomorphs in asexual reproduction. Mycologia 69: 1042–1045. Nimis, P. L. (1993) The Lichens of Italy. An Annotated Catalogue. Torino: Museo Regionale di Scienze Naturali, Monografie XII. Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. W. & Moore, D. M. (1992) The Lichen Flora of Great Britain and Ireland. London: Natural History Museum Publications. Rivas-Martı´nez, S. (1987) Mapa de las series de vegetación de Espan˜a 1:400 000 y memoria. Madrid: ICONA, Serie Técnica. Sanders, W. B. (1994) Role of lichen rhizomorphs in thallus propagation and substrate colonization. Cryptogamic Botany 4: 283–289. Sanders, W. B. & Rico, V. J. (1992) Lichenizing rhizomorphs and thallus development in the squamulose lichen Aspicilia crespiana Rico ined. (Lecanorales, Ascomycetes). Botanica Acta 105: 449–456. Santesson, R. (1993) The Lichens and Lichenicolous Fungi of Sweden and Norway. Lund: SBT-förlaget. Wagner, J. & Letrouit-Galinou, M.-A. (1988) Structure et ontogenèse du thalle squamuleux du lichen Endocarpon pusillum (Pyrénolichens, Verrucariacées). Canadian Journal of Botany 66: 2118–2127. White, F. J. & James, P. W. (1985) A new guide to microchemical techniques for the identification of lichen substances. British Lichen Society Bulletin 57 (Suppl.): 1–41. Wirth, V. (1995) Flechtenflora. Bestimmung und ökologische Kennzeichsnung der Flechten Südwestdeutschlands und angrenzender Gebiete. Stuttgart: Eugen Ulmer GmbH & Co. Accepted for publication 31 August 1998
ASPICILIA CRESPIANA, A NEW LICHEN SPECIES FROM SOUTHERN EUROPE Vı´ctor J. RICO*
Abstract: Aspicilia crespiana Rico is described as a new species from central Spain and Sardinia (Italy). In central Spain it is common and has a well-defined ecology. It is found at middle to high altitudes (from 400 to 1900 m), overgrowing mosses on exposed and horizontal to more or less vertical siliceous rocks. Closely related to the A. contorta (Hoffm.) Kremp. complex, A. crespiana differs in its muscicolous habit and thallus development, consisting of a network of lichenized squamules interlinked by fungal rhizomorphs. 1999 The British Lichen Society
Introduction In recent years a distinctive, muscicolous, squamulose to subfruticose Aspicilia s. lat., with strongly developed rhizomorphs, has been collected frequently in central Spain. Additional material from Sardinia (Italy) was sent to the author by Prof. P. L. Nimis (Trieste). The term ‘ rhizomorph ’ is used here in its broad sense, referring to linear hyphal aggregates with irregular structure, indeterminate growth, and with a generally exploratory function, as well as serving in attachment and propagation. Lichen rhizomorphs lack organized apices. Further information on the use of this term in lichenized fungi is given by Sanders (1994), and in non-lichenized fungi by Cairney et al. (1991). This taxon has been referred to as ‘ Aspicilia crespiana Rico ined. ’ in some publications (Nimis 1993; Sanders & Rico 1992). As this lichen apparently does not agree with any taxa in previously published papers on aspicilioid lichens it is described here as a new species. Materials and Methods Seventy-eight specimens of the new species, from seven provinces in Spain and one in Sardinia (Italy) were investigated (material deposited in MA, MAF and TSB). Freezing microtome sections (SLEE-MTC) and hand sections were mounted in water or in lactophenol-cotton-blue (Merck Rf. 13741). To study ascus structure, Lugol’s solution was added to sections and squash preparations after pre-treatment with KOH. A Nikon Optiphot-AFX II compound microscope was used for micrographs of sections and a Zeiss 47 50 52 dissecting microscope was used for habit photographs. Thirty-five selected specimens were analysed by TLC, following White & James (1985). Special care was taken with ascospore measurements and only well-developed and mature ascospores were analysed. These are more frequent in young and medium-sized apothecia of the studied material. The dimensions and length:breadth ratio (l:b) of ascospores are based on 93 *Departamento de Biologı´a Vegetal II, Facultad de Farmacia, Universidad Complutense, E–28040 Madrid, Spain. E-mail: [email protected] 0024–2829/99/020129+11 $30.00/0
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readings of several specimens of Aspicilia crespiana and on 40 readings of 4 specimens of A. contorta subsp. contorta and of A. contorta subsp. hoffmanniana (10 readings for each specimen cited below). Extreme values have been placed between brackets where they represented no more than 10% of the spores. For comparison of spore shape between taxa, I followed the terminology and corresponding l:b ratio defined by Bas (1969: 321–322) [globose (l:b of 1·0–1·05), subglobose (1·05–1·15), broadly ellipsoid (1·15–1·3), ellipsoid (1·3–1·6), elongate (1·6–2·0), cylindrical (2·0–3·0), bacilliform (>3·0)]. Besides the selected material of A. crespiana reported below, the following selected specimens were examined for comparison: Aspicilia contorta subsp. contorta: Spain: Burgos, Morentı´n, carretera de Castil de Peones a Villafranca-Montes de Oca, valle de Oca, monumento al General Mola, c. 950 m, guijarros calizos, 1996, G. Renobales (MAF). Madrid: Pelayos de la Presa, cruce del rı´o Alberche con la carretera comarcal 501, 550 m, 30TUK8969, granito con impregnaciones de polvo calizo, 1984, E. Barreno & V. J. Rico 32/1 (MAF); Perales de Tajun˜a, 650–780 m, 30TVK65, rocas calizas, 1980, E. Barreno & A. Merino (MAF). Navarra: Zudaire (Amezcoa Baja), 640 m, en la base de Quercus faginea, 1984, B. Aguirre (MAF). Aspicilia contorta subsp. hoffmanniana: Spain: Cáceres, Toril, Parque Natural de Monfragüe, Sierra del Serrejón, entre el Poblado del Embalse y La Herguijuela, Portilla del Tietar, 300 m, 30STK4716, cuarcitas, 1988, V. J. Rico 2088/2 (MAF). Madrid: Pelayos de la Presa, cruce del rı´o Alberche con la carratera comarcal 501, 550 m, 30TUK8969, granito, 1984, E. Barreno & V. J. Rico 55/2 (MAF); Hoyo de Manzanares, cerca de la urbanización La Berzosa, 980 m, 30TVK2196, granito, 1987, F. L. Redondo, Rico 1225/4 (MAF). Vizcaya: Ceánuri, Monte Larralde, 900 m, 30TWN2165, caliza compacta, 1985, M. Herrera & G. Renobales 1412 (BIO).
The Species Aspicilia crespiana Rico sp. nov. Aspicilia contortae affinis est, sed habito muscicola, thallo squamuloso vel subfruticuloso cum rhizomorphis, apotheciis distincte majoribus et sporis plerumque ellipsoidalibus differt. Thallus squamulosus vel subfruticulosus; crescit super Grimmia sp. in rupibus siliceus. Squamulae diametro usque ad 5 mm, imbricatae, dense congestae, apicem versus ascendentes vel lobatae, subtus ad rhizomorphis colligatae. Rhizomorphae bene evolutae, usque ad aliquot cm. longitudine, diametro usque ad 1·5 mm, irregulariter inaequaliter ramosae, algae destitutae; inter Grimmia sp. foliis et caulis et inter rupibus rimae penetrantes. Apothecia communis, primum cryptolecanorinia deinde lecideinia vel lecanorinia, immersa vel sessilia, usque ad 3·5 mm diametro; margine incrassato. Asci clavati, 2- vel 6-spori; tholis crassis, I. Ascosporae incolorae, simplices, plerumque ellipsoideae, (11–)15–32(–45)(8–)11–20(–31) ìm, cum tenuis pariete. Sine acido lichenoso. Typus: Spain, Madrid, San Martı´n de Valdeiglesias, road from Cadalso de los Vidrios to Pelayos de la Presa, km 3, Corcobada, 740 m, overgrowing Grimmia sp. on horizontal sun-exposed granitic rocks, 30TUK843668, 12 February 1988, V. J. Rico 1249/1 & M. A. Florido (MAF-Lich. 4221—holotypus; MA-Lichen 3274—isotypus).
(Figs 1–3) Thallus squamulose to subfruticose, consisting of commonly imbricate or dispersed lichenized squamules interconnected by abundant mycobiont rhizomorphs, forming large patches up to several decimetres diam.; squamules overgrowing saxicolous moss cushions (generally Grimmia species) and soil and rock in the vicinity, the rhizomorphs penetrating among these mosses and through the surrounding soil and rock. Squamules (Figs 1A, 2) up to 5 mm wide, highly variable. Mature squamules on mosses and soil, irregularly imbricate to contiguous, frequently fused to each other laterally, orbicular to elongate, generally ascending mainly at the tips, surface flat to concave or strongly convex, margins sinuate to lobulate, lobules rounded or commonly
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F. 1. Aspicilia crespiana. A, squamulose to subfruticose thallus growing on Grimmia sp. (holotype, MAF); B, lower side of thallus, showing rhizomorphs between mosses, rock fissures and soil; C, transverse section through mature lecanoroid apothecium in lactophenol-cotton-blue (holotype, MAF). Scales: A–B=2 mm; C=200 ìm.
verrucose to cylindrical and sometimes divided. Squamules of thallus margins on exposed, moss-free rocks, rare to frequent, sometimes areole-like, appressed, contiguous to scattered, orbicular to elongated,
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F. 2. Aspicilia crespiana (holotype, MAF). A, detail of thallus showing mature squamules overgrowing mosses; B, lateral view of squamules interlinked with rhizomorphs; C, detail of a subfruticose lichenized squamule (C1) with irregularly branched mycobiont rhizomorph (C2). Scale: 5 mm.
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margins mostly rounded, surface convex to commonly verrucose or bullate to ascending. Prothallic structures occasionally present on squamule margins, grey to white fan-like bands on moss leaves, dark strap-like bands on exposed rocks. Upper surface matt, white to grey or partly olivaceous to brownish, slightly roughened or smooth, frequently with white maculae consisting of accumulations of dead cortical cells; pycnidia or apothecial primordia also common as white to brown or black spots, without pseudocyphellae and pruina. Upper cortex 18–75(–90) ìm thick, paraplectenchymatous, of rounded cells with lumina 2·5–10(–12) ìm wide, brown-green pigmented at the surface; epinecral layer variable in thickness, consisting of dead mycobiont cell walls, without crystals or granules; tips of the ascending squamules also corticated below. Photobiont green, chlorococcoid, unicellular, cells 5–13 ìm diam.; arranged in a continuous layer or in a denticulate one when thallus squamules are thicker. Medulla loose to paraplectenchymatous, with pachydermatous hyphae, occasionally with grey granules, highly variable in thickness; strands of meso- to pachydermatous hyphae crossing this layer in a prosoplectenchymatous arrangement, extending from the squamules into the rhizomorphs. Lower surface of mature squamules white to ochraceous, without cortex or with a rudimentary one of 1, 2 or 3 rows of isodiametric cells, continuous with those of the rhizomorphs. Rhizomorphs (Figs 1B, 2) delicate, lacking algae, pale whitish or yellowish to pink and later darkly pigmented, terete to flattened, variable in length (up to several cm) and thickness (to 1·5 mm), branching extensively without any discernible pattern, interconnecting squamules terminally or laterally and giving rise to new ones. Rhizomorphs possess a central cord of densely packed meso- to pachy dermatous hyphae, in a prosoplectenchymatous arrangement. Surrounding the longitudinally arranged strands there is a rudimentary cortex of isodiametric cells (paraplectenchymatous) with or without pigmented walls. Distally from the squamules, the rhizomorphs eventually become fine, ecorticate and diffuse. Apothecia numerous, up to 3·5 mm wide, simple or composite, orbicular to compressed, immersed to elevated above the squamule or commonly sessile with constricted base, rarely slightly peltate, one to several per squamule, sometimes regenerating; cryptolecanorine or nearly urceolate at first, becoming lecanoroid (Fig. 1C) or lecideoid in larger fruits. Disc concave to flat, black. Margin distinct, later prominent, concolorous with thallus (lecanoroid) to brownish or black when thalline margin is reduced (lecideoid). Excipulum propium paraplectenchymatous in outer part, with brown-green to blackpigmented cell walls at the surface; highly variable in development and thickness, reduced in immersed apothecia to extended and rarely flabellate in large lecideoid fruits. Hypothecium colourless, composed of rounded cells, mostly reduced, with or without an algal layer underneath. Hymenium 105–160(–175) ìm high, conglutinated, colourless, without oil droplets; epithecium olivaceous to greenish, in young apothecia covered by dead cortical cells; subhymenium colourless. Paraphyses (Fig. 3B) branched and anastomosing, moniliform to submoniliform, with 1 to 9 rounded cells at the tips, up to 7 ìm thick at the olive-coloured apices, conglutinated. Asci (Fig. 3C) clavate, (75–)105–14020–40 ìm, 2- to 6-spored, mostly
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F. 3. Aspicilia crespiana. A, ascospores; A1 mature spore joined with the wall of a degenerating one; B, paraphyses; C, mature asci containing 2–6 spores; C1, ascus after treatment with I. Scale: 20 ìm.
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4-spored; apical apparatus thick, K/I; with a fuzzy, amyloid coat (Fig. 3C1). Ascospores (Fig. 3A) unicellular, colourless, thin-walled, rapidly degenerating at maturity, (11–)15–32(–45)(8–)11–20(–31) ìm, l:b ratio (1·03–)1·33–1·94(–2·42), l:b ratio mean value of 1·58, (globose) ellipsoid to elongate (cylindrical), mainly ellipsoid, rarely ovoid. Pycnidia common, more frequent at tips of squamules, immersed; ostiole greenish-brown to black; wall hyaline, rounded to convoluted; conidia 7–121 ìm, bacilliform, straight. Chemistry. No lichen substances detected by TLC; cortex and medulla K, C, KC, PD and I; pigment of epithecium and ostiolar area of pycnidia HNO3 + emerald green (Aspicilia-green). Etymology. This species is dedicated to Luis Crespı´ Jaume (Madrid, 1889–1963), who collected it during the nineteen twenties (cf. Crespı´ & González Bueno 1990). Distribution and habitat. Currently, A. crespiana is known from one locality in Sardinia (Italy) and from 14 localities in central Spain, where it is common and forms large, rather conspicuous patches. In A u vila (840 m alt.), Badajoz (725 m), Cáceres (400 m), Madrid (550–1300 m), Segovia (1130 m) and Toledo (580 m) provinces (Spain), it was collected in dry to subhumid situations from the mesomediterranean to supramediterranean belts of the Mediterranean Region (Rivas-Martı´nez 1987). It characteristically overgrows Grimmia species on siliceous rocks, mainly granite, usually in widely open situations amongst Quercus pyrenaica, Q. rotundifolia, Q. suber or cultivated Pinus pinea or P. sylvestris forests. In these localities a Cistus understorey is common. Aspicilia crespiana is found on sunny, horizontal to vertical rock surfaces, frequently with small amounts of sandy soil below the overgrown moss. The material from the Spanish province of Palencia (1900 m) is exceptional, having been collected in the subalpine belt of the Eurosiberian Region (Rivas-Martı´nez 1987), on exposed rock surfaces accompanied by few conspicuous gametophytes of an unidentified bryophyte. The most frequent lichens co-occurring with A. crespiana in central Spain are: Lecanora aff. muralis, Lepraria neglecta, Neofuscelia pulla, N. verruculifera, Xanthoparmelia tinctina and sometimes species of Cladonia and Lasallia pustulata. When growing near small rivers or raintracks, in slightly humid situations, Leptochidium albociliatum and Polychidium muscicola may also occur close to A. crespiana. Buellia badia and a taxon of the Rhizocarpon viriciatrum group are also common lichens growing on A. crespiana thalli. They together comprise a clearly differentiated community on mosses overgrowing horizontal or inclined, sun-exposed siliceous rocks. Remarks. The new species is easily recognized in the field by its primary muscicolous habit, and thallus development. Aspicilia crespiana is comprised of two interconnected structures, lichenized squamules and mycobiont rhizomorphs. The two types of structures can, under suitable conditions, give rise to each other. A detailed analysis of thallus development and variability, lichenizing rhizomorphs and substratum occupation of the new species is given by Sanders & Rico (1992). Similar types of structures have also been
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observed in other squamulose lichens, such as the terricolous Endocarpon pusillum (Malone 1977; Wagner & Letrouit-Galinou 1988) or the saxicolous Lecanora rhizinata (Barreno & Rico 1984; Sanders 1994). Terricolous species of Toninia frequently occur among mosses on soil, producing an extensive rhizomorphic system (Sanders 1994). The initial cryptolecanorine ascomatal structure, the ascus type with thick apical apparatus and amyloid gelatinous exoascal layer, moniliform paraphyses, olivaceous epithecium with HNO3 + reaction, and conidia morphology, confirms that this species belongs to the genus Aspicilia s. lat. (Clauzade & Roux 1984; Hafellner 1991; Purvis et al. 1992). The apothecial form, spore and conidial size, thickness of ascospore walls and number of spores per ascus, support its inclusion in Aspicilia subgen. Aspicilia (Clauzade & Roux 1984), and indicates that A. crespiana belongs to the A. calcarea (L.) Mudd group and is related to silicicolous forms of A. contorta (Hoffm.) Kremp. (Clauzade & Roux 1985; Esnault 1985; Ekman & Fröberg 1988; Hafellner 1991). Aspicilia contorta is a common lichen growing on calcareous rocks, limestone, mortar, siliceous rocks or rarely lignum of North Africa, Europe and North America (Clauzade & Roux 1985; Esnault 1985; Purvis et al. 1992). In Europe, it has a wide ecological amplitude and a correspondingly wide morphological variation (Nimis 1993). Several forms (Esnault 1985; Clauzade & Roux 1985) and intermediate specimens (Ekman & Fröberg 1988) have been described. For comparison, I follow Fröberg (1989) and Wirth (1995), who recognize two subspecies of A. contorta, A. contorta subsp. contorta and A. contorta subsp. hoffmanniana Ekman & Fröberg. Table 1 presents a comparison of the main taxonomic characters of A. crespiana and the two subspecies of A. contorta, based on recent, confirmed Spanish material (see Materials and Methods) and on bibliography. Aspicilia crespiana is clearly separated from A. contorta subsp. contorta and subsp. hoffmanniana by its muscicolous, squamulose to subfruticose thallus with strongly developed rhizomorphs. The apothecia and ascospore shape are also useful characters. Typically, the young apothecia of A. crespiana are similar to those (young and mature) of both subspecies of A. contorta, immersed, cryptolecanorine, with reduced excipulum propium and raised thallus margin. Mature apothecia of A. crespiana appear nearly urceolate, lecanoroid or lecideoid, and nearly all have a prominent margin. Moreover, a single mature apothecium of A. crespiana may have one part lecanoroid, with a prominent thalline margin and a reduced excipulum propium, and other parts lecideoid, with a prominent excipulum propium and a reduced or non-existent thalline margin. Sometimes, when apothecia are situated at the tips of ascending squamules or centred in convex ones, a short stalk can be distinguished. Despite the number of specimens examined, A. crespiana sporulated poorly, and only a limited number of mature and well-developed ascospores (93) could be measured. According to Bas (1969) the ascospores of A. crespiana are mainly ellipsoid in shape, and those of both subspecies of A. contorta are mainly broadly ellipsoid, as is inferred by comparing l:b ratio mean values (Table 1). Aspicilia contorta subsp. contorta and subsp. hoffmanniana occur together with the new species even in the same
0·5–2·5 mm diam., surface flat to convex, margins mainly rounded to lobulate
–
Up to 1·2 mm diam., immersed to subimmersed, cryptolecanorine to lecanoroid, margin later slightly raised, often pruinose
(20–)21–26(–30)(17–)18–22 (1·04–)1·09–1·33(–1·39) 1·20 (broadly ellipsoid)
Aspicilin ?*
Mainly on calcareous rocks, rarely on base enriched siliceous rocks or on lignum
N. C. & S. Europe‡
Areole/squamules
Rhizomorphs
Apothecia
Ascospore size (ìm) l:b ratio l:b ratio mean value
Chemistry
Habitat
Distribution (Europe)
N. C. & S. Europe‡
On calcareous or on siliceous rocks
Sometimes aspicilin ?*
(19–)20–30(–32)(15–)18–21(–25) (1·04–)1·1–1·55(–1·78) 1·28 (broadly ellipsoid)
Up to 1 mm diam., immersed to sessile, cryptolecanorine to lecanoroid, margin later raised to prominent, sometimes pruinose
–
0·1–1·8 mm diam., surface convex, margins mainly angled
Crustose, areolate, continuous to dispersed in thallus margins, grey to whitish, sometimes pruinose
subsp. hoffmanniana
C. Spain and Sardinia
Muscicolous and secondarily on siliceous rocks and soil in the vicinity
–
(11–)15–32(–45)(8–)11–20(–31) (1·03–)1·33–1·94(–2·42) 1·58(ellipsoid)
Up to 3·5 mm diam., immersed to sessile or slightly peltate, cryptolecanorine to lecanoroid or lecideoid, margin later prominent, never pruinose
+, Interconnecting squamules
Up to 5 mm diam., surface flat to concave or convex, margins with rounded, divided and cylindrical lobules, mainly ascending
Squamulose to subfruticose, continuous to dispersed in thallus margins, grey to whitish, never pruinose
A. crespiana
Aspicilia crespiana—Rico
*According to Ekman & Fröberg (1988). ‡According to Nimis (1993), Santesson (1993) and Wirth (1995).
Crustose, areolate to squamulose, dispersed in groups of scales, white to greyish, usually pruinose
subsp. contorta
Thallus
Characters
A. contorta
T 1. Main characters for distinguishing among Aspicilia contorta subsp. contorta, A. contorta subsp. hoffmanniana and A. crespiana
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locality, as seen in the studied material (Rico 32/1, Rico 55/2 and Rico 2968/1 respectively). However, it has a different habit in addition to the characters described above. Closely related to the A. contorta complex, and consequently with A. crespiana, are the fruticose, terricolous and vagrant species of the A. hispida Mereschk. agg. (Hafellner 1991). Morphologically, A. hispida is clearly different from the new species by its fruticose thallus, with cylindrical, pseudocyphellatae ramifications, lacking lichenized squamules and mycobiont rhizomorphs (Clauzade & Roux 1987). Additional selected specimens examined: Italy: Sardinia: Prov. Cagliari, Riserva Naturale di M. Arcosu, su muschi epilitici su trechite, 800 m, 1989, P. L. Nimis & M. Tretiach (TSB, MAF).—Spain: A u vila: Cebreros, en el km 1·4 de la carretera de Cebreros a El Hoyo de Pinares, ladera este del alto de El Castan˜ar, 840 m, 30TUK7680, sobre briófitos en granito, 1997, V. J. Rico 3329 (MAF). Badajoz: Garbayuela, Sierra de los Villares, a la derecha de la carretera desde Herrera del Duque, cerca de una cantera y antes del puerto de Los Carneros, 725 m, 30SUJ2829, sobre briófitos en cuarcitas y pizarras muy soleadas, 1996, V. J. Rico 3202 & M. A. Florido (MAF). Cáceres: junto a la Sierra de San Pedro, a 6 km por el camino que parte del suroeste de Malpartida de Cáceres, en las márgenes del rı´o Salor, 400 m, 29SQD1264, sobre briófitos en granito, 1992, F. J. Garcı´a González, Rico 2962 (MAF). Madrid: cerca de Hoyo de Manzanares, El Quemadillo, sobre musgos en pen˜as, 1926, L. Crespı´ (MA); San Martı´n de Valdeiglesias, embalse de San Juan: cerro Almodón, 650 m, 30TUK8273, sobre briófitos en granito, 1984, E. Barreno & V. J. Rico 114 (MAF); ibid., cerro Cabreruela, 690 m, 30TUK8573, sobre briófitos en granito, 1985, E. Barreno & V. J. Rico 334 (MAF); ibid., cerro de El Yelmo, 680 m, 30TUK8872, sobre briófitos en granito, 1991, V. J. Rico 2657 & M. A. Florido (MAF); Pelayos de la Presa, cruce del rı´o Alberche con la carretera comarcal 501, 550 m, 30TUK8969, granito, 1993, V. J. Rico 2968/1 (MAF); Cenicientos, pen˜a Cenicientos, 1253 m, 30TUK7259, 1986, J. M. Esteban, Manrique M34–40 (MAF); Robledo de Chavela, pico de El Almojón, 1175 m, 30TUK9581, sobre briófitos en granito, 1986, V. J. Rico 797 (MAF); San Lorenzo de El Escorial, Silla de Felipe II (bellow), on mosses over rock, exposed, 1991, W. B. Sanders (MAF); Lozoya, El Chaparral, 1300 m, 30TVL3634, sobre briófitos en gneis, 1986, V. J. Rico 744/1 & E. Manrique (MAF). Palencia: Resobada, pen˜a Curavacas, north face, 1900 m, 30TUN66, on siliceous conglomerate rock, no obvious bryophyte, 1992, W. B. Sanders (MAF). Segovia: San Ildefonso (La Granja), Parque de Santa Cecilia, cerca de la carretera de La Granja a Segovia, 1130 m, 30TVL1428, sobre briófitos en granito, 1998, V. J. Rico 3333/1 (MAF). Toledo: Nombela, en el km 13·3 de la carretera de Nombela a Pelahustán, 580 m, 30TUK6946, sobre briófitos en granito, 1997, V. J. Rico 3322 (MAF). I thank Dr C. Roux (Marseille) and Dr J. Esnault (Acigne) for their valuable comments on the specimens, Dr W. B. Sanders (San Francisco) for helping with the material and the photographs and for reviewing the manuscript, Dr G. Renobales (Vitoria) for the loan of Aspicilia contorta, Dr J. Pizarro (Madrid) for doing the line drawings, and Prof. P. L. Nimis (Trieste) for kindly sending me material from Sardinia. My dear friends Pepe Bella and Philip Mason are thanked also. R Barreno, E. & Rico, V. J. (1984) Sobre la biologı´a de los lı´quenes I. Anatomı´a, morfologı´a y estructuras vegetativas. Anales de Biologı´a (Murcia) 1: 161–195. Bas, C. (1969) Morphology and subdivision of Amanita and a monograph of its section Lepidella. Persoonia 5: 285–579. Cairney, J. W. G., Jennings, D. H. & Agerer, R. (1991) The nomenclature of fungal multi-hyphal linear aggregates. Cryptogamic Botany 2: 246–251. Clauzade, G. & Roux, C. (1984) Les genres Aspicilia Massal. et Bellemerea Hafellner et Roux. Bulletin de la Société Botanique du Centre-Ouest, Nouvelle série 15: 127–141. Clauzade, G. & Roux, C. (1985) Likenoj de Okcidenta Euˇropo. Ilustrita determinlibro. Bulletin de la Société Botanique du Centre-Ouest, Nouvelle série, numéro spécial 7-1985: 1–893.
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