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Two new species of Mycobilimbia from Spain
Lichenologist 35(1): 1-10 (2003) doi: 10.1006/lich.2002.0428
Two new species of My cobilimbia from Spain Francisco Jose SARRION, Gregorio ARAGON, Josef HAFELLNER, Victor J. RICO and Ana Rosa BURGAZ Abstract: Two new species of Mycobilimbia are described from Spain. Mycobilimbia olwacea has a minutely squamulose and olivaceous thalli and dark reddish brown apothecia, colourless inside; it has been found at middle to high altitudes (from 900 to 1750 m), growing on humid tree bases of Pmus tugra and lignum of Quercus pyrenaica. Mycobilimbia parvilobulosa is closely related to M. berengeriana, but differs markedly by its squamulose thalli, with crenate to subdigitiform margins, dark reddish brown excipulum and colourless epithecium. It grows on moss-covered old trunks of fagaceous trees. The new taxa are compared with M. berengeriana, M. hypnorum and M. sanguineoatra and their main distinguishing characters are tabulated. ( 2003 Published by Elsevier Science Ltd on behalf of The British Lichen Society. Key words: lichens, new species, Mycobilimbia, Spain.
species have been transferred from the Introduction There is much systematic and nomencla- genera Bacidia De Not., Biatora, Biatorina tural confusion surrounding the genera A. Massal., Bilimbia De Not., Lecidea Ach. Biatora Fr. and Mycobilimbia Rehm. Someor Toninia A. Massal. to Mycobilimbia, authors accept the genus Mycobilimbia mainly based on apothecium, ascus and (Vitikainen et al. 1997; Diederich & spore characters (Awasthi & Mathur 1987; Serusiaux 2000; Hafellner & Turk 2001; Hafellner 1984, 1989; Printzen 1995; Wirth Llimona & Hladun 2001) whilst others 1987, 1995). Nevertheless, Mycobilimbia is include it in Biatora (Purvis et al. 1992). still a heterogeneous and poorly known Moreover, Mycobilimbia as was 'originally' group, even in Europe where it is reprecircumscribed by Hafellner (1984) is sented by c. 15 species, and its nomenclature and taxonomy are undergoing change. heterogeneous. According to Hafellner (1984, 1989), the In recent years, some studies on both genus Mycobilimbia Rehm is separated from groups have been carried out, and several the genus Biatora Fr. by its well-developed thalli, ascospores with epispore and K/I + dark blue Mycobilimbia-type asci. Most F. J. Sarridn and A. R. Burgaz: Departamento de species of Biatora develop a crustose, effuse Biologia Vegetal I, Facultad de Biologia, Universidad Complutense, E-28040 Madrid, Spain. or granular thalli, ascospores without epiG. Aragon: Centro de Documentacion, Organismo spore and K/I + dark blue Biatora-type asci Autonomo Parques Nacionales (OAPN), Ministerio de (Purvis et al. 1992). Some years later, Medio Ambiente, c Gran Via de San Francisco, 4, Printzen (1995) based the separation of E-28005 Madrid, Spain. J. Hafellner: Institut fur Systematik Botanik, Karl- Biatora and Mycobilimbia on their differFranzens-Umversitat, Holteigasse 6, A-8010 Graz, ent apothecial ontogeny. The differences Austria. between Bacidia and Mycobilimbia are V. J. Rico: Departamento de Biologia Vegetal II, based mainly on thallus and apothecia Facultad de Farmacia, Universidad Complutense, Emorphology, apothecia ontogeny, shape, 28040 Madrid, Spain. 0024-2829 03 010001 + 10 S30.00 0
( 2003 Published by Elsevier Science Ltd on behalf of The British Lichen Society.
THE LICHENOLOGIST lumen and walls of excipulum hyphae, ascus structure and the ascospores (Ekman 1996). Toninia is mainly separated from Mycobilimbia by differences in the hamathecium of paraphyses, ascus structure and ascospores (Timdal 1991). The genus Myxobilimbia Hafellner (Hafellner & Tiirk 2001) has been recently described to accommodate species of the Bacidia sabuletorum group previously included in the genus Mycobilimbia. Following Ekman (1996), these species of the Bacidia sabuletorum group present a more developed ascus (wider walls), stronger paraphyses and a more developed excipulum than Mycobilimbia species. Differences in the colour of epithecium and hymenium, tholus type, ascospore size and shape, and habitat are also useful characters. During our studies on lichen biodiversity in Spain, many specimens belonging to the genus Mycobilimbia have been collected. Abundant material collected from two groups of populations could not be assigned to any known species. These specimens show a combination of characters which mainly accords with the current concept of Mycobilimbia in the broad sense of Hafellner (1989), but they are probably not closely related to the type of the genus (M. tetramera). However, since they are not conspecific with any other species of the genus or similar genera, they are described here as two new species of Mycobilimbia.
Material and Methods This study is based on 35 specimens, 20 of Mycobilimbia olivacea and 15 of Mycobilimbia parvilobulosa, of fresh material collected in several provinces in Spain (material deposited in MA, MACB, MAF and GZU). Freezing microtome sections and hand cut sections were mounted in water or in lactophenol cotton blue. To study ascus structure, Lugol's solution was added to sections and squash preparations after pre-treatment with KOH. Chemical analyses of selected specimens were carried out by thin layer chromatography (TLC), according to White & James (1985). All microscopy measurements were made on water mounts, and only well-developed mature ascospores were measured. Extreme values have been placed between brackets where they represent no more than 10% of the total number of measurements. The dimensions and length: width ratio (l:w) of ascospores are based on 120 readings of different specimens of Mycobilimbia olivacea and
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M. parvilobulosa. Current mycological terminology is used and generally follows Kirk et al. (2001), and phytoclimatic terminology is according to RivasMartinez (1987). The following selected specimens were examined for comparison and compilation of Table 1: Mycobilimbia berengeriana (A. Massal.) Hafellner & V. Wirth. Austria: Salzburg: Hohe Tauern, Hafer Gruppe, Lungau, Murtal, Rotgtilden, Aufstieg zum Unteren Rotgiilden-See, 1370-1710 m, Hochstaudenfluren, 1981, J. Poelt, H. Mayrhofer & R. Tiirk (GZU). Tirol: Otztaler Alpen, Otztal, 3 km SSE Obergurgl, Abstieg von der Wetterstation Barenhoppet (am Riicken der Hohen Mut) ins Gaisbergtal, 24502600 m, MTB 9132/3, substrat: moose, 1993, W. Obermayer 2761 (GZU).—Canada: Ontario: Thunder Bay District, Ouimet, at Ouimet Canyon, 48°45'N, 88°35'W, on a dry rock table outcrop, and in the adjoining ravine leading into the canyon, at base of Thuja, 1965, /. M. Brodo 6108, R. Cain & B. Denison (GZU).—Greenland: Disko: Umbebung von Godhavn, Unteres Blasedal NE Godhavn, 1982, J. Poelt & H. Ullrich (GZU).—Hungary: prope pagum Malyinka, in declivibus montis Czakoko, alt. c. 570 m.s.m., 1938, F. Foriss exs. 47 (GZU).—Italy: Sardinia: Prov. Nuoro, Barbagia Seulo, Road SeuiLanusei, M. Arcueri, close to Cantoniera Arcueri, 950-1100 m, Quercus ilex wood, limestone, siliceous rocks, 9°21'E, 39°49'N, 1975, J. Poelt (GZU).--Spain: Albacete: Alcaraz, Sierra de Alcaraz, rio del Barrancazo, 30SWH4969, 1350 m, en la base musgosa de Quercus faginea, 1997, G. Aragon 1116/97 & I. Martinez (MA). Jaen: La Iruela, Sierra de Cazorla, rio Guadalquivir, 30SWH0702, 850 m, en la base de Quercus rotimdifolia, 1996, G. Aragon 0733/96 & V. J. Rico (MAF); Siles, Sierra de Segura, Las Acebeas, 30SWH3641, 1350 m, sobre briofitos, en talud calcareo, 1997, G. Aragon 1662/97 & I. Martinez (MA). Navarra: Isaba, Larra, Km 22 despues del tiinel, 1650 m, en restos vegetales, 1995, J. Etayo (MA); Lanz, paredes calcareas siguiendo el torrente bajo el hayedo, 750 m, fisuras en calizas verticales, 1994, J. Etayo (MA).—Sweden: Lyckscle Lappmark: Tarna, Umfors 19 20/7, at 600-700 m.s.m., on decaying vegetation, earth and moss in the wood, A. H. Magnusson exs. 206 (GZU).—USA: Colorado: Hinsdale Co., along trail to west of Rio Grande Reservoir, spruce forest area, 37°46'30"NT, 107°21'30"W, 2870-3000 m, on mosses, 1984, T. H. Nash 25715 (GZU). Michigan: Isle Royale National Park, North side of Washington Harbor just W of Beaver Island, wet drainage to shore with Thuja, spruce and birch, 1984, C. M. Wetmore 52958 (GZU). Oregon: Benton County, 8-1 km N of Corvallis, 44°39'N, 123°19'W, 450 m.s.m., on bark and mosses on trunks of Quercus garryana, 1973, L. H. Pike exs. 428 (GZU). Mycobilimbia hypnorum (Libert) Kalb & Hafellner. Austria: Steiermark: Eisenerzer Alpen, Reichenstein NW von Trofaiach, zwischen der Krumpalm und dem Krumpensee, c. 1420 m, 47°29'30"N, 14°56'30"E, MTB 8555/2 nieder Kalkblocke in einem subalpinen Weiderasen, 1996, J. Hafellner 39699 & I. Martinez (GZU); ibid., auf Peltigera elisabethae, 1996, J. Hafellner
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Two new species of Mycobilimbia—Sarrion et al.
39883 fr /. Martinez (GZU).—Italy: Friuli-Venezia Giulia: Prov. Udinc, Carnics Alps, Monte Fleone (Raudenspitz), slopes NE obove Giogo Verania (Ofer Joch), c. 2100 m, 46°38'30"N, 12°44'50"E, paleozoic micaschists over bn'ophvtes and plants remnants, 1996, J. Hafellner 39183'(GZU).—Spain: Albacete: Riopar, Sierra del Calar del Mundo, entre Fuente de la Pedorrilla y Cueva de los Chorros, 30SWH4956, 1350 m, en grietas calizas, sobre briofitos, 1996, G. Aragon 267/96, A. Herrero & I. Martinez (MA). Jam: Siles, Sierra de Segura, Las Acebeas, 30SWH3641, 1350 m, en taludes musgosos, 1996, G. Aragon 899/96 & I. Martinez (MAF); Siles, Sierra de Segura, Las Acebeas, 30SWH3641, 1350 m, en fisuras calcareas, sobre briofitos, 1997, G. Aragon \Tibl97 & 1. Martinez (MA); Homos, Sierra de Cazorla, Sierra de las Lagunillas, cerca del embalse del Tranco, 30SWH1720, 830 m, sobre rocas cubiertas de briofitos, 1997, G. Aragon 2242/97 (MAF). Malaga: Parauta, estnbaciones de la Sierra de las Nieves, 30SUF1559, 970 m, en la base de encina, 1995, G. Aragon & I. Martinez (MA). Kavarra: Larra, base del monte Arias, La Contienda, 1750 m, briofitos calcicolas, 1995, J. A. Azpilicueta, J. Etayo & W. Mutter (MA). Soria: Santa Maria de las Hoyas, Sierra de Nafria, canon del rio Lobos, 30TVM9125, 1050 m, en grietas humedas, sobre briofitos, 1996, G. Aragon 1300/96 & I. Martinez (MA). Toledo: Hontanar, Montes de Toledo, arroyo del Gatillo, 30SUJ7384, 1100 m, en la base musgosa de Quercus pyrenaica, 1995, G. Aragon 859/95, A. Herrero fr /. Martinez (MA). Zamora: Cobreros, valle de Sanabria, Sotillo de Sanabria, arroyo de las Truchas, 29TPG8662, 1230 m, sobre briofitos, en la base de Querais pyrenaica, 1997, G. Aragon 651/97 & /. Martinez (MA).
The Species Mycobilimbia olivacea Aragon, Sarrion & Hafellner sp. nov. Thallus granulosus vel squamulosus, olivaceus; squamulae 0-2-0-7(-l) mm in diametro, dense congestae, = imbricatae. Apothecia communis, lecideinia, sessilia, 0-2-0-8( 1) mm in diametro, disci atrorubro-brunnea, epruinosi. Asci typo Mycobilimbia. Ascosporae hyalinae, simplices, ellipsoideae vel angustae ellipsoideae, (9-) 11-15(-17) x 5-6-5(-7-5) (jm, episporio laevi. Typus: Spain, Jaen, Quesada, Sierra de Cazorla, arroyo del Arteson, 30SVG9988, 1450 m, pine forest, on the base of Pimts nigra, 6 April 1998, G. Aragon 0819 98 & I. Martinez (MA 12766—holotypus; MA— isotypus).
(Figs 1 & 2) Thallus corticolous, crustose, continuous, thick, minutely squamulose, dark green to dark olive-brown; consisting of commonly continuous, ± imbricate squamules, 0-2-
0-7(-l) mm diameter, up to 0-1 mm high, surface flat to convex, margins sinuate to finely lobulate. Upper cortex thin, up to 10 fim, consisting of 1 or 2 rows of ± rounded cells with lumina 4-5-9 um wide, brown to brown-orange pigmented at the surface. Photobiont green, chlorococcoid, cells 5-9 um diameter, arranged in a continuous layer, 45-60 um thick. Ascomata apothecia, numerous, lecideoid to biatorine, 0-2-0-8(-l) mm diam., simple, ± orbicular to compressed, commonly sessile, with ± constricted base; disc flat to convex, dark reddish brown. Excipulum ± prosoplectenchymatous in outer part, well-developed, 48-55 um wide, with reddish brown pigmented cell walls at the surface, cells in outer part with lumina 6-8 um wide, those in inner part with ± enlarged lumina, up to 13 um. Epithecium ochraceous-brown, 5-12 um high. Hymenium (45-)50-70(-80) um high, colourless. Hypothecium 50-70(-75) um high, colourless, containing photobiont cells in the inner part. Paraphyses 1-5-2-5 jam wide, sparsely branched and anastomosing, apices 3-4(-5) um, becoming clavate and with an orange-brown apical cap. Asci clavate, 8-spored, apical apparatus thick, K/I + dark blue, containing a more densely amyloid tubular pore, of Mycobilimbia-type. Ascospores biseriate, colourless, unicellular, with smooth epispore, (9-) 11-15 (-17) x 5-6-5(-7-5) um, l:w ratio (l-67-)2-2-66 (-2-96), l:w ratio mean value of 2-36, elongate to cylindrical or ovoid. Conidiomata not found. Chemistry. No lichen substances detected by TLC; cortex, K - , C - , KC - , Pd - . Etymology. The specific epithet refers to the olivaceous colour of the thallus. Habitat and distribution. Mycobilimbia olivacea was found in humid to subhumid meso- and supramediterranean belts (RivasMartinez 1987) growing in mature forests, lining the fissures of tree bases in dense Pinus nigra formations and on lignum of Quercus pyrenaica in southern parts of
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FIG. 1. Thallus morphology in Mycobilimbia. A, M. olivacea (holotvpe); B & C, M. parvilobulosa {Aragoti 686/95, MA). Scales: A & B = O5 mm; 6 = 0-25 mm.
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Two new species of Mycobilimbia—Sarrion et al.
FIG. 2. Mycobilimbia olivacea {Sarrion 1187, MACB). A, habitus; B, section through a mature apothecium and thallus; C, ascospores. Scales: A=0-2 mm; B = 50 um; C = 5 um.
THE LICHENOLOGIST the Iberian Peninsula. The Pinus nigra forests cover extensive areas in supra- and oromediterranean belts growing on rock outcrops on the calcareous mountains of the eastern Peninsula. There are also wellpreserved forests in 'Sierra de Segura' (Jaen province) and 'Sierra de Alcaraz' (Albacete province) which have a good cover (80%), and are located in deep valleys covering vast areas of potentially Quercus faginea forests mixed with maples (Acer granatense) and holly (Ilex aquifolium). In 'Serrania de Cuenca' (Cuenca province), black pine occurs on some of the less disturbed dolines where the forests are well-structured. On Pinus nigra, M. olivacea is found with Cladonia comocraea, C. ochrochlora, Hypocenomyce anthracophila, H. scalaris, Micarea prasina, Placynthiella icmalea and Trapeliopsis flexuosa. The pyrenean oak forests of 'Sierra Madrona' (Ciudad Real province) grow on siliceous substrata in subhumid meso- and supramediterranean belts mixed with whitebeams (Sorbus torminalis), maples (Acermonspessulanum), strawberry tree {Arbutus unedo) and wild cherry (Prunus avium). The stands are clearly age-structured and there is no evidence of recent human disturbance. In this case, M. olivacea is frequently found together with Cladonia chlorophaea, C. ramulosa, Evernia prunastri, Hypogymnia tubulosa, Parmelia saxatilis, P. sulcata, Parmelina tiliacea, Placynthiella icmalea, Platismatia glauca and Trapeliopsis flexuosa. Remarks. The ascocarp ontogeny and thin-walled spores with smooth epispore of the new species are comparable to the genus Biatora (Prinzen 1995). However, the welldeveloped thalli, and the asci with a Kl + dark blue reaction and containing a tubular pore are characteristic of the genus Mycobilimbia. The relationships between Mycobilimbia species with this combination of characters and Biatora need further investigation. Mycobilimbia olivacea is easily recognized by its minutely squamulose dark green to olive brown thallus and dark reddish brown apothecia. Other Mycobilimbia species (Table 1) with a similar morphology
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and widespread in the Iberian Peninsula are M. berengeriana and M. hypnorum. Thallus morphology and substratum ecology of M. olivacea resemble a Hypocenomyce species, although there are important differences of paraphyses, shape and size of ascospores, ascus structure and chemistry. Additional specimens examined. Spain: Albacete: Paterna del Madera, Sierra de Alcaraz, Calar de la Osera, 30SWH5167, 1550 m, sobre Pinus nigra, 1997, G. Aragon 870/97 & I. Martinez (MA); Bienservida, Sierra de Alcaraz, Padron de Bienservida, 30SWH3961, 1600 m, sobre Pinus nigra, 1997, G. Aragon 1389/97, A. Herrero & I. Martinez (MA); Bogarra, Sierra de Alcaraz, cerca de Las Espineras, 30SWH4966, 1580 m, sobre Pinus nigra, 1996, G. Aragon 2030/96, A. Herrero & A. Pujol (MA); Riopar, Sierra del Calar del Mundo, cntre Fuente de la Pedorrilla y Cueva de los Chorros, 30SWH4956, 1350 m, sobre Pinus nigra, 1996, G. Aragon 0290/96, A Herrero & I. Martinez (MA). Ciudad Real: Fuencaliente: robledal de 'El Abuelo', 30SUH7956, 950 m, sobre un basidiocarpo seco de hongo lignicola, en tronco muerto de Quercus pyrenaica, 1997, A. R. Burgaz, I. Martinez & F. J. Sarrion 1187 (MACB); ibid., Sierra de Quintana, umbria de Burcio del Pino, 30SUH9351, 1010 m, en tocon de Quercus pyrenaica, 1998, F. J. Sarrion 2589 (MACB), 2120 (MACB, GZU). Cuenca: Cuenca, Serrania de Cuenca: pista que va dcsdc Las Majadas a Una, 30TWK8758, 1400 m, sobre Pinus nigra, 1998, G. Aragon & I. Martinez 2124 (MA); ibid., Las Torcas, 30TWK8834, 1300 m, sobre Pinus nigra, 1998, G. Aragon & I. Martinez 2724 (MA); Buenache de la Sierra, Serrania de Cuenca, Rodenales, 30TWK8942, 1350 m, sobre Pinus nigra, 1998, G. Aragon & I. Martinez 2574 (MA); La Cierva, Serrania de Cuenca, Dehesa de los Palancares, Las Torcas, 30TWK9132, 1200 m, sobre Pinus nigra, 1998, G. Aragon & I. Martinez 2675 (MA). Jaen: Siles, Sierra de Segura, Las Acebeas, 30SWH3641, 1350 m, sobre Pinus nigra, 1996, G. Aragon 0871/96 & I. Martinez (MAF); ibid., 1997, G. Aragon 1660/97 & I. Martinez (MA); Segura de la Sierra, Sierra de Segura, rio Madera, 30SWH3331, 1200 m, sobre Pinus nigra, 1996, G. Aragon 1067/96 & /. Martinez (MA); Pozo Alcon, Sierra del Pozo, subida al cerro Cabanas, 30SWG0484,1750 m, sobre Pinus nigra, 1997, G. Aragon 2305/97 (MA).
Mycobilimbia parvilobulosa Sarrion, Aragon & Hafellner sp. nov. Thallus granulosus vel squamulosus, pallide viridis vel glaucus; squamulae 0-2-0-8 mm in diametro, ± imbricatae, apicem parvilobulosae. Apothecia communis, lecideinia, sessilia, 0-3-1 mm in diametro; disci atrobrunnei, epruinosi. Asci typo Mycobilimbia. Ascosporae simplices, cllipticae vel angustae, (9-5-) 11 -14(—16-5) x 4-6 urn, episporco laevi vel aspero.
Apices to 3 \m\ Pale brown to yellowish brown sometimes with scattered blue (K+green) Dark reddish brown
Apices to 5 6 ^m Colourless
Dark reddish brown
Epispore finely warted On bryophytes over old fagaceous tree trunks
Colourless
Epispore smooth On acid bark and lignum
Dark reddish brown
Epispore finely warted On bryophytes over calcareous rocks, soils and old fagaceous tree trunks
Yellowish brown
Dark reddish brown, sometimes with scattered blue (K + green) granules
Epispore finely warted On bryophytes over calcareous rocks and old fagaceous tree trunks
Epithecium
Hypothecium
Ascosporcs Habitat
*Based on: Hafellncr (1989); Purvis ct al. (1992) (see Lecidea berengcriana and L. hypnorum); Wirth (1995); fresh material, fBascd on Purvis et al. (1992) (see Lecidea sanguineoatra).
Ochraceous-brown
Epispore smooth On bryophytes over acid rocks or bark
Colourless, with scattered blue (K+green) granules
Colourless, without blue granules
Colourless, with scattered blue (K+green) granules
Colourless, sometimes with scattered blue (K+green) granules in the inner part Apices to 5-6 (im
Hymenium
Apices to 3-4 \xm
Dark reddish brown
Dark reddish brown
Colourless; outer edge reddish brown Colourless, without blue granules
Dark reddish brown
Dark reddish brown
Excipulum
Apices to 3 (im Pale brown to yellowish brown sometimes with scattered blue (K+ green) granules
Convex, pale to dark brown
Flat to convex, dark brown or blackish
Flat to convex, dark reddish brown
Flat to convex (sometimes tuberculate), dark brown to black
Flat to convex, dark brown or blackish
Apothecia
Paraphyses
Thin, effuse, membranous, whitish or greenish grey
Thick, squamulose, 0-2-0-8 mm diam., margins crenate to subdigitiform, whitish to greenish grey
Thick, squamulose, 0-2-0-7(-l) mm diam., dark green to dark olive brown
Thin, effuse, membranous, whitish or greenish grey
M. sanguineoatra^
M. pan'ilobulosa
Al. olivacea
Al. hypnorum*
Thick, granular warts, 0-04-0-1 (-0-35) mm diam., whitish to greenish grey
Al. bcretigcriaua*
Thallus
Character
TABLE 1. Pri)icipal characters for distinguishing Mycobilimbia berengcriana, M. hypnorum, M. olivacea, M. parvilobulosa and M. sanguineoatra
rt M
2"
o
O
n ft'
ft
5
13 ft
to o o u
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FIG. 3. Mycobilimbia parvilobulosa (holotype, Sanion 1236, MACB). A, habitus; B, partial section of hymenium; C, ascospores. Scales: A = O 2 m m ; B = 50um; C = 5|.im.
lose to minutely squamulose, whitish to greenish grey; granules 0-07-0-17 mm wide, developing into minute squamules; squamules 0-2-0-8 wide x 0-15-0-2 mm high, surface flat, margin crenate to sub(Figs 1 & 3) digitiform, weakly corticate. Upper cortex Thallus corticolous, crustose, widespread thin, up to 15 um, surrounded by a to continuous and imbricate, thick, granu- closely compacted layer of hyphae of 1-3 um
Typus: Spain, Ciudad Real, Solana del Pino, Banos de las Tinosas, on the bark of Qitercus faginea subsp. broteroi, 4 February 1997, A. R. Burgas, I. Martinez <2" F. J. Sarnon 1236 (MACB 82733—holotypus; MA— isotypus).
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Two new species of Mycobilimbia—Sanion et al.
diam. Photobiont green, chlorococcoid, cells 8-15 |im diam., arranged in a continuous layer 55-75 jam thick. Medulla of loose hyphae, 2-4 |im diam., projecting into the substratum below the squamules. Ascomata apothecia, numerous, lecideoid to biatorine, 0-3-1 mm diam., simple, ± orbicular to compressed, with ± constricted base; disc flat to convex, dark brown; margin thick and darker at first, later excluded and apothecia sometimes finally tuberculate. Excipulum ± prosoplectenchymatous, of radiating hyphae, with thick, reddish brown walls, darker in the outer part; cells of inner part in contact with the hypothecium with lumina 6-11 x 1-2 |am, outer cells swollen and darker with lumina 7-9 x 4 6 |am. Epithecium colourless, with sparse orange gelatinous granules in the surface. Hymenium (50-)60-70(-75) urn high, colourless. Hypothecium (55-)60-75(-80) um high, dark reddish brown. Paraphyses 2-2-5 |jm wide, mostly simple, apices 4-5 urn, becoming clavate or capitate. Asci clavate, 8-spored, apical apparatus of Mycobilimbia-type. Ascospores uni-biseriate, colourless, unicellular, with finely warted epispore, (9-5-)l 1 —14(—16-5) x 4-6 |am, l:w ratio (l-8-)2-2-66(-3), l:w ratio mean value of 2-4, ellipsoid to fusiform. Conidiomata not found. Chemistry. No lichen substances detected by TLC; cortex K — or K ± yellow. Etymology. The specific epithet refers to small (Latin, parvus) lobules of the thallus. Habitat and distribution. Mycobilimbia parvilobulosa has been found on old fagaceous tree trunks ( ± near the base), in hyperhumid to subhumid situations of the meso- and supramediterranean belts (RivasMartinez 1987) of the southern Iberian Peninsula. It usually overgrows bryophytes and occurs in old woodland sites together with Collema subnigrescens, Fuscopannaria ignobilis, F. mediterranean Leptogium lichenoides, L. saturninum, Lobaria pulmonaria, L. scrobiculata, Nephroma laevigatum, Peltigera collina and P. horizontalis. It was collected in 'Sierra Madrona' (Ciudad Real province)
and 'Montes de Toledo' (Toledo province) from Quercus faginea subsp. broteroi growing in narrow valleys on siliceous substrata. In 'Sierra de Segura' (Jaen province) and 'Sierra de Alcaraz' (Albacete province) the species grows on Q. faginea subsp. faginea and Q. ilex subsp. ballota, which cover deep limestone valleys and steep ravines. The species was also detected on mossy trunks of old Q. faginea subsp. faginea in relict pinsapo fir forests (Abies pinsapo) of southern Spain (Cadiz province). Remarks. Mycobilimbia parvilobulosa is clearly distinguished from M. berengeriana, M. hypnorum and M. sanguineoatra by its granulose to squamulose thallus with crenate to subdigitiform margins and a thallus type similar to that of Phyllopsora species (Hafellner 1989). The absence of pigmented granules (K+green) in the hymenium and hypothecium of M. parvilobulosa is also a useful character (see Table 1). Furthermore, there are differences between the species in the size and colour of the apices of paraphyses. The paraphyses of M. hypnorum and M. sanguineoatra are slender and have less swollen apices (up to 2-5 (im) than M. berengeriana and M. parvilobulosa, with apices to 5-6 (j.m. In addition, the apices of paraphyses are brown-walled in M. berengeriana and colourless in the other four species. Additional specimens examined. Spain: Albacete: Alcaraz, Sierra de Alcaraz, rio del Barrancazo, 30SWH4969, 1350 m, en la base de Quercus faginea, 1997, G. Aragdn 1116/97 & /. Martinez (MA). Cadiz: Grazalema, umbria de la Sierra del Pinar, 1100 m, sobre Quercus Jaginea subsp. faginea, 1997, F. J. Sarrion 753 (MACB, GZU). Ciudad Real: Fuencaliente, Sierra Madrona: Jerumbrosa, 30SUH8551, 800 m, sobre Quercus faginea subsp. broteroi, 1997, F. J. Sarrion 1652 (MACB); ibid., arroyo del Robledo de las Hoyas, 30SUH8256, 800 m, sobre Quercus faginea subsp. broteroi, 1997, A. R. Burgaz, I. Martinez & F. J. Sarrion 2469 (MACB). Jaen: La Iruela, Sierra de Segura, rio Borosa, 30SWH1305, 900 m, sobre Quercus ilex subsp. ballota, 1995, G. Aragdn 683/95, 686/95, 688/95 & /. Martinez (MA, MAF); Santiago-Pontones, Sierra de Segura, rio Aguamulas, cerca del cortijo del Mulon, 30SWH1711, 800 m, en la base de Quercus ilex subsp. ballota, 1995, G. Aragdn 282/95 & I. Martinez (MA). Toledo: Hontanar, Montes de Toledo, arroyo del Gatillo, 30SUJ7384, sobre Quercus pyrenaica, 1995, G. Aragdn 859/95, A. Herrero & /. Martinez (MA); Los Navalucillos, Montes de Toledo, Las Becerras, arroyo
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del Chorro, 30SUJ5978, 950 m, en base musgosa de Quercus pyrenaica, 1995, G. Aragon 129/95, A. Herrero & I. Martinez (MA). We are very grateful to Christian Printzen and the anonymous referee for their valuable comments. This study was supported by the Spanish 'Junta de Comunidades de Castilla-La Mancha, Consejeria de Agricultura y Medio Ambiente, Direction General del Medio Natural'. Dr Sarrion received financial support from the Universidad Complutense de Madrid to visit the Institut fur Systematik Botanik, Karl FranzensUniversitat, Graz, Austria.
REFERENCES
Awasthi, D. D. & Mathur, R. (1987) Species of the lichen genera Bacidia, Badimia, Fellhanera and Mycobilimbia from India. Proceedings of the Indian Academy of Sciences {Plant Sciences) 97: 481-503. Diederich, P. & Serusiaux, E. (2000) The Lichens and Lichenicolous Fungi of Belgium and Luxemburg. Luxembourg: Musee National d'Histoire Naturelle. Ekman, S. (1996) The corticolous and lignicolous species of Bacidia and Bacidina in North America. Opera Botanka 127: 1-148. Hafellner, J. (1984) Studien in Richtung einer natiirlicheren Gliederung der Sammelfamilien Lecanoraceae und Lecideaceae. Nova Hedwigia Beihefte 79: 241-371. Hafellner, J. (1989) Die europaischen MycobilimbiaArten eine erste Ubersicht (lichenisierte Ascomyceten, Lecanorales). Herzogia 8: 53-89.
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Hafellner, J. & Turk, R. (2001) Die lichenisierten Pilze Osterreichs—eine Checkliste der bisher nachgewiesenen Arten mit Verbreitungsangaben. Stapfia 76: 1-167. Kirk, P, M., Cannon, P. F., David, J. C. & Stalpers, J. A. (2001) Ainsivorth & Bisby's Dictionary of the Fungi. 9th Edn. Wallingford: CAB International. Llimona, X. & Hladun, N. (2001) Checklist of the lichens and lichenicolous fungi of the Iberian Peninsula and Balearic Islands. Bocconea 14: 5-581. Printzen, C. (1995) Die Flechtengattung Biatora in Europa. Bibliotheca Lichenologica 60: 1-275. Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. W. & Moore, D. M. (eds) (1992) The Lichen Flora of Great Britain and Ireland. London: Natural History Museum Publications. Rivas-Martinez, S. (1987) Mapa de las series de vegetacion de Espaha 1: 400.000y memoria. Madrid: ICONA, Serie Tecmca. Timdal, E. (1991) A monograph of the genus Toninia {Lecideaceae, Ascomycetes). Opera Botanica 110: 1-137. Vitikainen, O., Ahti, T., Kuusinen, M., Lommi, S. & Ulvinen, T. (1997) Checklist of lichens and allied fungi of Finland. Norrlinia 6: 1-123. White, F. J. & James, P. W. (1985) A new guide to microchemical techniques for the identification of lichen substances. British Lichen Society Bulletin 57, Supplement: 1-41. Wirth, V. (1987) Die Flechten Baden-Wurttembcrgs. Stuttgart: Ulmer. Wirth, V. (1995) Flechtenflora. Stuttgart: Eugen Ulmer GmbH & Co.
Accepted for publication 28 October 2002
Two new species of My cobilimbia from Spain Francisco Jose SARRION, Gregorio ARAGON, Josef HAFELLNER, Victor J. RICO and Ana Rosa BURGAZ Abstract: Two new species of Mycobilimbia are described from Spain. Mycobilimbia olwacea has a minutely squamulose and olivaceous thalli and dark reddish brown apothecia, colourless inside; it has been found at middle to high altitudes (from 900 to 1750 m), growing on humid tree bases of Pmus tugra and lignum of Quercus pyrenaica. Mycobilimbia parvilobulosa is closely related to M. berengeriana, but differs markedly by its squamulose thalli, with crenate to subdigitiform margins, dark reddish brown excipulum and colourless epithecium. It grows on moss-covered old trunks of fagaceous trees. The new taxa are compared with M. berengeriana, M. hypnorum and M. sanguineoatra and their main distinguishing characters are tabulated. ( 2003 Published by Elsevier Science Ltd on behalf of The British Lichen Society. Key words: lichens, new species, Mycobilimbia, Spain.
species have been transferred from the Introduction There is much systematic and nomencla- genera Bacidia De Not., Biatora, Biatorina tural confusion surrounding the genera A. Massal., Bilimbia De Not., Lecidea Ach. Biatora Fr. and Mycobilimbia Rehm. Someor Toninia A. Massal. to Mycobilimbia, authors accept the genus Mycobilimbia mainly based on apothecium, ascus and (Vitikainen et al. 1997; Diederich & spore characters (Awasthi & Mathur 1987; Serusiaux 2000; Hafellner & Turk 2001; Hafellner 1984, 1989; Printzen 1995; Wirth Llimona & Hladun 2001) whilst others 1987, 1995). Nevertheless, Mycobilimbia is include it in Biatora (Purvis et al. 1992). still a heterogeneous and poorly known Moreover, Mycobilimbia as was 'originally' group, even in Europe where it is reprecircumscribed by Hafellner (1984) is sented by c. 15 species, and its nomenclature and taxonomy are undergoing change. heterogeneous. According to Hafellner (1984, 1989), the In recent years, some studies on both genus Mycobilimbia Rehm is separated from groups have been carried out, and several the genus Biatora Fr. by its well-developed thalli, ascospores with epispore and K/I + dark blue Mycobilimbia-type asci. Most F. J. Sarridn and A. R. Burgaz: Departamento de species of Biatora develop a crustose, effuse Biologia Vegetal I, Facultad de Biologia, Universidad Complutense, E-28040 Madrid, Spain. or granular thalli, ascospores without epiG. Aragon: Centro de Documentacion, Organismo spore and K/I + dark blue Biatora-type asci Autonomo Parques Nacionales (OAPN), Ministerio de (Purvis et al. 1992). Some years later, Medio Ambiente, c Gran Via de San Francisco, 4, Printzen (1995) based the separation of E-28005 Madrid, Spain. J. Hafellner: Institut fur Systematik Botanik, Karl- Biatora and Mycobilimbia on their differFranzens-Umversitat, Holteigasse 6, A-8010 Graz, ent apothecial ontogeny. The differences Austria. between Bacidia and Mycobilimbia are V. J. Rico: Departamento de Biologia Vegetal II, based mainly on thallus and apothecia Facultad de Farmacia, Universidad Complutense, Emorphology, apothecia ontogeny, shape, 28040 Madrid, Spain. 0024-2829 03 010001 + 10 S30.00 0
( 2003 Published by Elsevier Science Ltd on behalf of The British Lichen Society.
THE LICHENOLOGIST lumen and walls of excipulum hyphae, ascus structure and the ascospores (Ekman 1996). Toninia is mainly separated from Mycobilimbia by differences in the hamathecium of paraphyses, ascus structure and ascospores (Timdal 1991). The genus Myxobilimbia Hafellner (Hafellner & Tiirk 2001) has been recently described to accommodate species of the Bacidia sabuletorum group previously included in the genus Mycobilimbia. Following Ekman (1996), these species of the Bacidia sabuletorum group present a more developed ascus (wider walls), stronger paraphyses and a more developed excipulum than Mycobilimbia species. Differences in the colour of epithecium and hymenium, tholus type, ascospore size and shape, and habitat are also useful characters. During our studies on lichen biodiversity in Spain, many specimens belonging to the genus Mycobilimbia have been collected. Abundant material collected from two groups of populations could not be assigned to any known species. These specimens show a combination of characters which mainly accords with the current concept of Mycobilimbia in the broad sense of Hafellner (1989), but they are probably not closely related to the type of the genus (M. tetramera). However, since they are not conspecific with any other species of the genus or similar genera, they are described here as two new species of Mycobilimbia.
Material and Methods This study is based on 35 specimens, 20 of Mycobilimbia olivacea and 15 of Mycobilimbia parvilobulosa, of fresh material collected in several provinces in Spain (material deposited in MA, MACB, MAF and GZU). Freezing microtome sections and hand cut sections were mounted in water or in lactophenol cotton blue. To study ascus structure, Lugol's solution was added to sections and squash preparations after pre-treatment with KOH. Chemical analyses of selected specimens were carried out by thin layer chromatography (TLC), according to White & James (1985). All microscopy measurements were made on water mounts, and only well-developed mature ascospores were measured. Extreme values have been placed between brackets where they represent no more than 10% of the total number of measurements. The dimensions and length: width ratio (l:w) of ascospores are based on 120 readings of different specimens of Mycobilimbia olivacea and
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M. parvilobulosa. Current mycological terminology is used and generally follows Kirk et al. (2001), and phytoclimatic terminology is according to RivasMartinez (1987). The following selected specimens were examined for comparison and compilation of Table 1: Mycobilimbia berengeriana (A. Massal.) Hafellner & V. Wirth. Austria: Salzburg: Hohe Tauern, Hafer Gruppe, Lungau, Murtal, Rotgtilden, Aufstieg zum Unteren Rotgiilden-See, 1370-1710 m, Hochstaudenfluren, 1981, J. Poelt, H. Mayrhofer & R. Tiirk (GZU). Tirol: Otztaler Alpen, Otztal, 3 km SSE Obergurgl, Abstieg von der Wetterstation Barenhoppet (am Riicken der Hohen Mut) ins Gaisbergtal, 24502600 m, MTB 9132/3, substrat: moose, 1993, W. Obermayer 2761 (GZU).—Canada: Ontario: Thunder Bay District, Ouimet, at Ouimet Canyon, 48°45'N, 88°35'W, on a dry rock table outcrop, and in the adjoining ravine leading into the canyon, at base of Thuja, 1965, /. M. Brodo 6108, R. Cain & B. Denison (GZU).—Greenland: Disko: Umbebung von Godhavn, Unteres Blasedal NE Godhavn, 1982, J. Poelt & H. Ullrich (GZU).—Hungary: prope pagum Malyinka, in declivibus montis Czakoko, alt. c. 570 m.s.m., 1938, F. Foriss exs. 47 (GZU).—Italy: Sardinia: Prov. Nuoro, Barbagia Seulo, Road SeuiLanusei, M. Arcueri, close to Cantoniera Arcueri, 950-1100 m, Quercus ilex wood, limestone, siliceous rocks, 9°21'E, 39°49'N, 1975, J. Poelt (GZU).--Spain: Albacete: Alcaraz, Sierra de Alcaraz, rio del Barrancazo, 30SWH4969, 1350 m, en la base musgosa de Quercus faginea, 1997, G. Aragon 1116/97 & I. Martinez (MA). Jaen: La Iruela, Sierra de Cazorla, rio Guadalquivir, 30SWH0702, 850 m, en la base de Quercus rotimdifolia, 1996, G. Aragon 0733/96 & V. J. Rico (MAF); Siles, Sierra de Segura, Las Acebeas, 30SWH3641, 1350 m, sobre briofitos, en talud calcareo, 1997, G. Aragon 1662/97 & I. Martinez (MA). Navarra: Isaba, Larra, Km 22 despues del tiinel, 1650 m, en restos vegetales, 1995, J. Etayo (MA); Lanz, paredes calcareas siguiendo el torrente bajo el hayedo, 750 m, fisuras en calizas verticales, 1994, J. Etayo (MA).—Sweden: Lyckscle Lappmark: Tarna, Umfors 19 20/7, at 600-700 m.s.m., on decaying vegetation, earth and moss in the wood, A. H. Magnusson exs. 206 (GZU).—USA: Colorado: Hinsdale Co., along trail to west of Rio Grande Reservoir, spruce forest area, 37°46'30"NT, 107°21'30"W, 2870-3000 m, on mosses, 1984, T. H. Nash 25715 (GZU). Michigan: Isle Royale National Park, North side of Washington Harbor just W of Beaver Island, wet drainage to shore with Thuja, spruce and birch, 1984, C. M. Wetmore 52958 (GZU). Oregon: Benton County, 8-1 km N of Corvallis, 44°39'N, 123°19'W, 450 m.s.m., on bark and mosses on trunks of Quercus garryana, 1973, L. H. Pike exs. 428 (GZU). Mycobilimbia hypnorum (Libert) Kalb & Hafellner. Austria: Steiermark: Eisenerzer Alpen, Reichenstein NW von Trofaiach, zwischen der Krumpalm und dem Krumpensee, c. 1420 m, 47°29'30"N, 14°56'30"E, MTB 8555/2 nieder Kalkblocke in einem subalpinen Weiderasen, 1996, J. Hafellner 39699 & I. Martinez (GZU); ibid., auf Peltigera elisabethae, 1996, J. Hafellner
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Two new species of Mycobilimbia—Sarrion et al.
39883 fr /. Martinez (GZU).—Italy: Friuli-Venezia Giulia: Prov. Udinc, Carnics Alps, Monte Fleone (Raudenspitz), slopes NE obove Giogo Verania (Ofer Joch), c. 2100 m, 46°38'30"N, 12°44'50"E, paleozoic micaschists over bn'ophvtes and plants remnants, 1996, J. Hafellner 39183'(GZU).—Spain: Albacete: Riopar, Sierra del Calar del Mundo, entre Fuente de la Pedorrilla y Cueva de los Chorros, 30SWH4956, 1350 m, en grietas calizas, sobre briofitos, 1996, G. Aragon 267/96, A. Herrero & I. Martinez (MA). Jam: Siles, Sierra de Segura, Las Acebeas, 30SWH3641, 1350 m, en taludes musgosos, 1996, G. Aragon 899/96 & I. Martinez (MAF); Siles, Sierra de Segura, Las Acebeas, 30SWH3641, 1350 m, en fisuras calcareas, sobre briofitos, 1997, G. Aragon \Tibl97 & 1. Martinez (MA); Homos, Sierra de Cazorla, Sierra de las Lagunillas, cerca del embalse del Tranco, 30SWH1720, 830 m, sobre rocas cubiertas de briofitos, 1997, G. Aragon 2242/97 (MAF). Malaga: Parauta, estnbaciones de la Sierra de las Nieves, 30SUF1559, 970 m, en la base de encina, 1995, G. Aragon & I. Martinez (MA). Kavarra: Larra, base del monte Arias, La Contienda, 1750 m, briofitos calcicolas, 1995, J. A. Azpilicueta, J. Etayo & W. Mutter (MA). Soria: Santa Maria de las Hoyas, Sierra de Nafria, canon del rio Lobos, 30TVM9125, 1050 m, en grietas humedas, sobre briofitos, 1996, G. Aragon 1300/96 & I. Martinez (MA). Toledo: Hontanar, Montes de Toledo, arroyo del Gatillo, 30SUJ7384, 1100 m, en la base musgosa de Quercus pyrenaica, 1995, G. Aragon 859/95, A. Herrero fr /. Martinez (MA). Zamora: Cobreros, valle de Sanabria, Sotillo de Sanabria, arroyo de las Truchas, 29TPG8662, 1230 m, sobre briofitos, en la base de Querais pyrenaica, 1997, G. Aragon 651/97 & /. Martinez (MA).
The Species Mycobilimbia olivacea Aragon, Sarrion & Hafellner sp. nov. Thallus granulosus vel squamulosus, olivaceus; squamulae 0-2-0-7(-l) mm in diametro, dense congestae, = imbricatae. Apothecia communis, lecideinia, sessilia, 0-2-0-8( 1) mm in diametro, disci atrorubro-brunnea, epruinosi. Asci typo Mycobilimbia. Ascosporae hyalinae, simplices, ellipsoideae vel angustae ellipsoideae, (9-) 11-15(-17) x 5-6-5(-7-5) (jm, episporio laevi. Typus: Spain, Jaen, Quesada, Sierra de Cazorla, arroyo del Arteson, 30SVG9988, 1450 m, pine forest, on the base of Pimts nigra, 6 April 1998, G. Aragon 0819 98 & I. Martinez (MA 12766—holotypus; MA— isotypus).
(Figs 1 & 2) Thallus corticolous, crustose, continuous, thick, minutely squamulose, dark green to dark olive-brown; consisting of commonly continuous, ± imbricate squamules, 0-2-
0-7(-l) mm diameter, up to 0-1 mm high, surface flat to convex, margins sinuate to finely lobulate. Upper cortex thin, up to 10 fim, consisting of 1 or 2 rows of ± rounded cells with lumina 4-5-9 um wide, brown to brown-orange pigmented at the surface. Photobiont green, chlorococcoid, cells 5-9 um diameter, arranged in a continuous layer, 45-60 um thick. Ascomata apothecia, numerous, lecideoid to biatorine, 0-2-0-8(-l) mm diam., simple, ± orbicular to compressed, commonly sessile, with ± constricted base; disc flat to convex, dark reddish brown. Excipulum ± prosoplectenchymatous in outer part, well-developed, 48-55 um wide, with reddish brown pigmented cell walls at the surface, cells in outer part with lumina 6-8 um wide, those in inner part with ± enlarged lumina, up to 13 um. Epithecium ochraceous-brown, 5-12 um high. Hymenium (45-)50-70(-80) um high, colourless. Hypothecium 50-70(-75) um high, colourless, containing photobiont cells in the inner part. Paraphyses 1-5-2-5 jam wide, sparsely branched and anastomosing, apices 3-4(-5) um, becoming clavate and with an orange-brown apical cap. Asci clavate, 8-spored, apical apparatus thick, K/I + dark blue, containing a more densely amyloid tubular pore, of Mycobilimbia-type. Ascospores biseriate, colourless, unicellular, with smooth epispore, (9-) 11-15 (-17) x 5-6-5(-7-5) um, l:w ratio (l-67-)2-2-66 (-2-96), l:w ratio mean value of 2-36, elongate to cylindrical or ovoid. Conidiomata not found. Chemistry. No lichen substances detected by TLC; cortex, K - , C - , KC - , Pd - . Etymology. The specific epithet refers to the olivaceous colour of the thallus. Habitat and distribution. Mycobilimbia olivacea was found in humid to subhumid meso- and supramediterranean belts (RivasMartinez 1987) growing in mature forests, lining the fissures of tree bases in dense Pinus nigra formations and on lignum of Quercus pyrenaica in southern parts of
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FIG. 1. Thallus morphology in Mycobilimbia. A, M. olivacea (holotvpe); B & C, M. parvilobulosa {Aragoti 686/95, MA). Scales: A & B = O5 mm; 6 = 0-25 mm.
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Two new species of Mycobilimbia—Sarrion et al.
FIG. 2. Mycobilimbia olivacea {Sarrion 1187, MACB). A, habitus; B, section through a mature apothecium and thallus; C, ascospores. Scales: A=0-2 mm; B = 50 um; C = 5 um.
THE LICHENOLOGIST the Iberian Peninsula. The Pinus nigra forests cover extensive areas in supra- and oromediterranean belts growing on rock outcrops on the calcareous mountains of the eastern Peninsula. There are also wellpreserved forests in 'Sierra de Segura' (Jaen province) and 'Sierra de Alcaraz' (Albacete province) which have a good cover (80%), and are located in deep valleys covering vast areas of potentially Quercus faginea forests mixed with maples (Acer granatense) and holly (Ilex aquifolium). In 'Serrania de Cuenca' (Cuenca province), black pine occurs on some of the less disturbed dolines where the forests are well-structured. On Pinus nigra, M. olivacea is found with Cladonia comocraea, C. ochrochlora, Hypocenomyce anthracophila, H. scalaris, Micarea prasina, Placynthiella icmalea and Trapeliopsis flexuosa. The pyrenean oak forests of 'Sierra Madrona' (Ciudad Real province) grow on siliceous substrata in subhumid meso- and supramediterranean belts mixed with whitebeams (Sorbus torminalis), maples (Acermonspessulanum), strawberry tree {Arbutus unedo) and wild cherry (Prunus avium). The stands are clearly age-structured and there is no evidence of recent human disturbance. In this case, M. olivacea is frequently found together with Cladonia chlorophaea, C. ramulosa, Evernia prunastri, Hypogymnia tubulosa, Parmelia saxatilis, P. sulcata, Parmelina tiliacea, Placynthiella icmalea, Platismatia glauca and Trapeliopsis flexuosa. Remarks. The ascocarp ontogeny and thin-walled spores with smooth epispore of the new species are comparable to the genus Biatora (Prinzen 1995). However, the welldeveloped thalli, and the asci with a Kl + dark blue reaction and containing a tubular pore are characteristic of the genus Mycobilimbia. The relationships between Mycobilimbia species with this combination of characters and Biatora need further investigation. Mycobilimbia olivacea is easily recognized by its minutely squamulose dark green to olive brown thallus and dark reddish brown apothecia. Other Mycobilimbia species (Table 1) with a similar morphology
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and widespread in the Iberian Peninsula are M. berengeriana and M. hypnorum. Thallus morphology and substratum ecology of M. olivacea resemble a Hypocenomyce species, although there are important differences of paraphyses, shape and size of ascospores, ascus structure and chemistry. Additional specimens examined. Spain: Albacete: Paterna del Madera, Sierra de Alcaraz, Calar de la Osera, 30SWH5167, 1550 m, sobre Pinus nigra, 1997, G. Aragon 870/97 & I. Martinez (MA); Bienservida, Sierra de Alcaraz, Padron de Bienservida, 30SWH3961, 1600 m, sobre Pinus nigra, 1997, G. Aragon 1389/97, A. Herrero & I. Martinez (MA); Bogarra, Sierra de Alcaraz, cerca de Las Espineras, 30SWH4966, 1580 m, sobre Pinus nigra, 1996, G. Aragon 2030/96, A. Herrero & A. Pujol (MA); Riopar, Sierra del Calar del Mundo, cntre Fuente de la Pedorrilla y Cueva de los Chorros, 30SWH4956, 1350 m, sobre Pinus nigra, 1996, G. Aragon 0290/96, A Herrero & I. Martinez (MA). Ciudad Real: Fuencaliente: robledal de 'El Abuelo', 30SUH7956, 950 m, sobre un basidiocarpo seco de hongo lignicola, en tronco muerto de Quercus pyrenaica, 1997, A. R. Burgaz, I. Martinez & F. J. Sarrion 1187 (MACB); ibid., Sierra de Quintana, umbria de Burcio del Pino, 30SUH9351, 1010 m, en tocon de Quercus pyrenaica, 1998, F. J. Sarrion 2589 (MACB), 2120 (MACB, GZU). Cuenca: Cuenca, Serrania de Cuenca: pista que va dcsdc Las Majadas a Una, 30TWK8758, 1400 m, sobre Pinus nigra, 1998, G. Aragon & I. Martinez 2124 (MA); ibid., Las Torcas, 30TWK8834, 1300 m, sobre Pinus nigra, 1998, G. Aragon & I. Martinez 2724 (MA); Buenache de la Sierra, Serrania de Cuenca, Rodenales, 30TWK8942, 1350 m, sobre Pinus nigra, 1998, G. Aragon & I. Martinez 2574 (MA); La Cierva, Serrania de Cuenca, Dehesa de los Palancares, Las Torcas, 30TWK9132, 1200 m, sobre Pinus nigra, 1998, G. Aragon & I. Martinez 2675 (MA). Jaen: Siles, Sierra de Segura, Las Acebeas, 30SWH3641, 1350 m, sobre Pinus nigra, 1996, G. Aragon 0871/96 & I. Martinez (MAF); ibid., 1997, G. Aragon 1660/97 & I. Martinez (MA); Segura de la Sierra, Sierra de Segura, rio Madera, 30SWH3331, 1200 m, sobre Pinus nigra, 1996, G. Aragon 1067/96 & /. Martinez (MA); Pozo Alcon, Sierra del Pozo, subida al cerro Cabanas, 30SWG0484,1750 m, sobre Pinus nigra, 1997, G. Aragon 2305/97 (MA).
Mycobilimbia parvilobulosa Sarrion, Aragon & Hafellner sp. nov. Thallus granulosus vel squamulosus, pallide viridis vel glaucus; squamulae 0-2-0-8 mm in diametro, ± imbricatae, apicem parvilobulosae. Apothecia communis, lecideinia, sessilia, 0-3-1 mm in diametro; disci atrobrunnei, epruinosi. Asci typo Mycobilimbia. Ascosporae simplices, cllipticae vel angustae, (9-5-) 11 -14(—16-5) x 4-6 urn, episporco laevi vel aspero.
Apices to 3 \m\ Pale brown to yellowish brown sometimes with scattered blue (K+green) Dark reddish brown
Apices to 5 6 ^m Colourless
Dark reddish brown
Epispore finely warted On bryophytes over old fagaceous tree trunks
Colourless
Epispore smooth On acid bark and lignum
Dark reddish brown
Epispore finely warted On bryophytes over calcareous rocks, soils and old fagaceous tree trunks
Yellowish brown
Dark reddish brown, sometimes with scattered blue (K + green) granules
Epispore finely warted On bryophytes over calcareous rocks and old fagaceous tree trunks
Epithecium
Hypothecium
Ascosporcs Habitat
*Based on: Hafellncr (1989); Purvis ct al. (1992) (see Lecidea berengcriana and L. hypnorum); Wirth (1995); fresh material, fBascd on Purvis et al. (1992) (see Lecidea sanguineoatra).
Ochraceous-brown
Epispore smooth On bryophytes over acid rocks or bark
Colourless, with scattered blue (K+green) granules
Colourless, without blue granules
Colourless, with scattered blue (K+green) granules
Colourless, sometimes with scattered blue (K+green) granules in the inner part Apices to 5-6 (im
Hymenium
Apices to 3-4 \xm
Dark reddish brown
Dark reddish brown
Colourless; outer edge reddish brown Colourless, without blue granules
Dark reddish brown
Dark reddish brown
Excipulum
Apices to 3 (im Pale brown to yellowish brown sometimes with scattered blue (K+ green) granules
Convex, pale to dark brown
Flat to convex, dark brown or blackish
Flat to convex, dark reddish brown
Flat to convex (sometimes tuberculate), dark brown to black
Flat to convex, dark brown or blackish
Apothecia
Paraphyses
Thin, effuse, membranous, whitish or greenish grey
Thick, squamulose, 0-2-0-8 mm diam., margins crenate to subdigitiform, whitish to greenish grey
Thick, squamulose, 0-2-0-7(-l) mm diam., dark green to dark olive brown
Thin, effuse, membranous, whitish or greenish grey
M. sanguineoatra^
M. pan'ilobulosa
Al. olivacea
Al. hypnorum*
Thick, granular warts, 0-04-0-1 (-0-35) mm diam., whitish to greenish grey
Al. bcretigcriaua*
Thallus
Character
TABLE 1. Pri)icipal characters for distinguishing Mycobilimbia berengcriana, M. hypnorum, M. olivacea, M. parvilobulosa and M. sanguineoatra
rt M
2"
o
O
n ft'
ft
5
13 ft
to o o u
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FIG. 3. Mycobilimbia parvilobulosa (holotype, Sanion 1236, MACB). A, habitus; B, partial section of hymenium; C, ascospores. Scales: A = O 2 m m ; B = 50um; C = 5|.im.
lose to minutely squamulose, whitish to greenish grey; granules 0-07-0-17 mm wide, developing into minute squamules; squamules 0-2-0-8 wide x 0-15-0-2 mm high, surface flat, margin crenate to sub(Figs 1 & 3) digitiform, weakly corticate. Upper cortex Thallus corticolous, crustose, widespread thin, up to 15 um, surrounded by a to continuous and imbricate, thick, granu- closely compacted layer of hyphae of 1-3 um
Typus: Spain, Ciudad Real, Solana del Pino, Banos de las Tinosas, on the bark of Qitercus faginea subsp. broteroi, 4 February 1997, A. R. Burgas, I. Martinez <2" F. J. Sarnon 1236 (MACB 82733—holotypus; MA— isotypus).
2003
Two new species of Mycobilimbia—Sanion et al.
diam. Photobiont green, chlorococcoid, cells 8-15 |im diam., arranged in a continuous layer 55-75 jam thick. Medulla of loose hyphae, 2-4 |im diam., projecting into the substratum below the squamules. Ascomata apothecia, numerous, lecideoid to biatorine, 0-3-1 mm diam., simple, ± orbicular to compressed, with ± constricted base; disc flat to convex, dark brown; margin thick and darker at first, later excluded and apothecia sometimes finally tuberculate. Excipulum ± prosoplectenchymatous, of radiating hyphae, with thick, reddish brown walls, darker in the outer part; cells of inner part in contact with the hypothecium with lumina 6-11 x 1-2 |am, outer cells swollen and darker with lumina 7-9 x 4 6 |am. Epithecium colourless, with sparse orange gelatinous granules in the surface. Hymenium (50-)60-70(-75) urn high, colourless. Hypothecium (55-)60-75(-80) um high, dark reddish brown. Paraphyses 2-2-5 |jm wide, mostly simple, apices 4-5 urn, becoming clavate or capitate. Asci clavate, 8-spored, apical apparatus of Mycobilimbia-type. Ascospores uni-biseriate, colourless, unicellular, with finely warted epispore, (9-5-)l 1 —14(—16-5) x 4-6 |am, l:w ratio (l-8-)2-2-66(-3), l:w ratio mean value of 2-4, ellipsoid to fusiform. Conidiomata not found. Chemistry. No lichen substances detected by TLC; cortex K — or K ± yellow. Etymology. The specific epithet refers to small (Latin, parvus) lobules of the thallus. Habitat and distribution. Mycobilimbia parvilobulosa has been found on old fagaceous tree trunks ( ± near the base), in hyperhumid to subhumid situations of the meso- and supramediterranean belts (RivasMartinez 1987) of the southern Iberian Peninsula. It usually overgrows bryophytes and occurs in old woodland sites together with Collema subnigrescens, Fuscopannaria ignobilis, F. mediterranean Leptogium lichenoides, L. saturninum, Lobaria pulmonaria, L. scrobiculata, Nephroma laevigatum, Peltigera collina and P. horizontalis. It was collected in 'Sierra Madrona' (Ciudad Real province)
and 'Montes de Toledo' (Toledo province) from Quercus faginea subsp. broteroi growing in narrow valleys on siliceous substrata. In 'Sierra de Segura' (Jaen province) and 'Sierra de Alcaraz' (Albacete province) the species grows on Q. faginea subsp. faginea and Q. ilex subsp. ballota, which cover deep limestone valleys and steep ravines. The species was also detected on mossy trunks of old Q. faginea subsp. faginea in relict pinsapo fir forests (Abies pinsapo) of southern Spain (Cadiz province). Remarks. Mycobilimbia parvilobulosa is clearly distinguished from M. berengeriana, M. hypnorum and M. sanguineoatra by its granulose to squamulose thallus with crenate to subdigitiform margins and a thallus type similar to that of Phyllopsora species (Hafellner 1989). The absence of pigmented granules (K+green) in the hymenium and hypothecium of M. parvilobulosa is also a useful character (see Table 1). Furthermore, there are differences between the species in the size and colour of the apices of paraphyses. The paraphyses of M. hypnorum and M. sanguineoatra are slender and have less swollen apices (up to 2-5 (im) than M. berengeriana and M. parvilobulosa, with apices to 5-6 (j.m. In addition, the apices of paraphyses are brown-walled in M. berengeriana and colourless in the other four species. Additional specimens examined. Spain: Albacete: Alcaraz, Sierra de Alcaraz, rio del Barrancazo, 30SWH4969, 1350 m, en la base de Quercus faginea, 1997, G. Aragdn 1116/97 & /. Martinez (MA). Cadiz: Grazalema, umbria de la Sierra del Pinar, 1100 m, sobre Quercus Jaginea subsp. faginea, 1997, F. J. Sarrion 753 (MACB, GZU). Ciudad Real: Fuencaliente, Sierra Madrona: Jerumbrosa, 30SUH8551, 800 m, sobre Quercus faginea subsp. broteroi, 1997, F. J. Sarrion 1652 (MACB); ibid., arroyo del Robledo de las Hoyas, 30SUH8256, 800 m, sobre Quercus faginea subsp. broteroi, 1997, A. R. Burgaz, I. Martinez & F. J. Sarrion 2469 (MACB). Jaen: La Iruela, Sierra de Segura, rio Borosa, 30SWH1305, 900 m, sobre Quercus ilex subsp. ballota, 1995, G. Aragdn 683/95, 686/95, 688/95 & /. Martinez (MA, MAF); Santiago-Pontones, Sierra de Segura, rio Aguamulas, cerca del cortijo del Mulon, 30SWH1711, 800 m, en la base de Quercus ilex subsp. ballota, 1995, G. Aragdn 282/95 & I. Martinez (MA). Toledo: Hontanar, Montes de Toledo, arroyo del Gatillo, 30SUJ7384, sobre Quercus pyrenaica, 1995, G. Aragdn 859/95, A. Herrero & /. Martinez (MA); Los Navalucillos, Montes de Toledo, Las Becerras, arroyo
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del Chorro, 30SUJ5978, 950 m, en base musgosa de Quercus pyrenaica, 1995, G. Aragon 129/95, A. Herrero & I. Martinez (MA). We are very grateful to Christian Printzen and the anonymous referee for their valuable comments. This study was supported by the Spanish 'Junta de Comunidades de Castilla-La Mancha, Consejeria de Agricultura y Medio Ambiente, Direction General del Medio Natural'. Dr Sarrion received financial support from the Universidad Complutense de Madrid to visit the Institut fur Systematik Botanik, Karl FranzensUniversitat, Graz, Austria.
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Hafellner, J. & Turk, R. (2001) Die lichenisierten Pilze Osterreichs—eine Checkliste der bisher nachgewiesenen Arten mit Verbreitungsangaben. Stapfia 76: 1-167. Kirk, P, M., Cannon, P. F., David, J. C. & Stalpers, J. A. (2001) Ainsivorth & Bisby's Dictionary of the Fungi. 9th Edn. Wallingford: CAB International. Llimona, X. & Hladun, N. (2001) Checklist of the lichens and lichenicolous fungi of the Iberian Peninsula and Balearic Islands. Bocconea 14: 5-581. Printzen, C. (1995) Die Flechtengattung Biatora in Europa. Bibliotheca Lichenologica 60: 1-275. Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. W. & Moore, D. M. (eds) (1992) The Lichen Flora of Great Britain and Ireland. London: Natural History Museum Publications. Rivas-Martinez, S. (1987) Mapa de las series de vegetacion de Espaha 1: 400.000y memoria. Madrid: ICONA, Serie Tecmca. Timdal, E. (1991) A monograph of the genus Toninia {Lecideaceae, Ascomycetes). Opera Botanica 110: 1-137. Vitikainen, O., Ahti, T., Kuusinen, M., Lommi, S. & Ulvinen, T. (1997) Checklist of lichens and allied fungi of Finland. Norrlinia 6: 1-123. White, F. J. & James, P. W. (1985) A new guide to microchemical techniques for the identification of lichen substances. British Lichen Society Bulletin 57, Supplement: 1-41. Wirth, V. (1987) Die Flechten Baden-Wurttembcrgs. Stuttgart: Ulmer. Wirth, V. (1995) Flechtenflora. Stuttgart: Eugen Ulmer GmbH & Co.
Accepted for publication 28 October 2002