Ramalina carminae (Ascomycota: Ramalinaceae), una especie nueva de Europa.
The genus Ramalina Ach. is a cosmopolitan and very species-rich lichen genus. It comprises c. 200 species according to Fletcher et al. (2009). It is regarded as one of the taxonomically most difficult genera of macrolichens because most of its species are quite variable in their morphology, probably due to environmental modifications, and many of them show also variability in their secondary metabolites. There have been made several approaches to the knowledge of this genus, considering not only the morphology of the species, but also their anatomy and secondary metabolites, being the later essential in the comprehesion of this genus (Krog & James 1977, Krog & 0sthagen 1980, Stevens 1987, Bartsch 1992, Kashiwadani & Kalb 1993, Blanchon et al. 1996, Bannister & Blanchon 2003, Aptroot & Bungartz 2007, Aptroot & Schumm 2008).
The high intraspecific chemical variability of some of its species is not correlated with a morphological variation. In the past, this led to the description of a number of infraspecific and even specific taxa, that were subsequently treated as chemical strains or chemotypes. In Europe and America key studies were made in the Ramalina farinacea group (Culberson 1966, Hawksworth 1968, Krog & James 1977, Bowler & Rundel 1978, Krog & 0sthagen 1980) and the R. siliquosa group (Culberson 1967, Culberson & Culberson 1967, Hawksworth 1976, Sheard & James 1976, S0chting 1976, Krog & James 1977, Santesson 1984, Clauzade & Roux 1985).
In the Iberian Peninsula this genus has been widely studied, from a morphological and a chemical point of view and many contributions to the Iberian Lichenological Flora are presented (Arroyo & Manrique 1988, Arroyo & Manrique 1989, Arroyo et al. 1991, Arroyo 1993, Arroyo et al. 1995, Alvarez et al. 2001).
The morphological variability which characterizes some species of this genus, such as the Ramalina polymorpha group, has caused many problems in order to understand it. From Acharius (1810) to the present, numerous studies have dealt with the taxonomy of this group which includes Ramalina polymorpha (Ach.) Ach., R. capitata (Ach.) Nyl., R. protecta H. Magn., R. strepsilis (Ach.) Zahlbr. and R. digitellata Nyl. In some cases this taxa can be distinguished morphologically, however intermediate forms often appear, which has led to a permanent discussion over its taxonomic position. Some authors (Krog & James 1977, Krog & 0sthagen 1980) recognized a single species Ramalina polymorpha (Ach.) Ach. with different morphotypes, but this synthetic approach has not been accepted by other authors who have recognized in the group two species: Ramalina polymorpha (Ach.) Ach. and R. capitata (Ach.) Nyl. (Santesson 1984, Sancho 1988, Nimis & Poelt 1987).
This two species were recognized in the chemical and morphological study of the Ramalina polymorpha group in Spain (Arroyo et al. 1991) which was based on 891 specimens. This study also suggested to treat the different morphotypes observed among R. capitata, which is a very polymorphic species, especially in the Mediterranean area, as varieties. Nimis (1993) gave an infraspecific range to this morphotypes: R. capitata (Ach.) Nyl. var. capitata,, R. capitata var. protecta (Magnusson) Nimis and R. capitata var. digitellata (Nyl.) Nimis.
In the present work, mainly based on material collected by the authors, we describe a new species from Portugal, Spain and Sardinia (Italy). The new species is associated to the different varieties which, according to Nimis (1993) we accept among R. capitata. We will refere to these varieties as Ramalina capitata complex.
MATERIALS AND METHODS
The first material studied (67 specimens) was collected by the authors on several field trips in the Tiermes archaeological site, at Montejo de Tiermes (SW of the province of Soria, Spain). We have done the Ramalina genus revision (year 2007) in different national and international herbaria and we found two specimens from Sardinia (Italy) and three from Portugal, which are identical in its morphology as well as in its chemical composition to those presented in this work. 140 specimens collected by the authors and other different collectors between 1967 and 1990 in different Spanish locations were provisionally included into the Ramalina capitata complex as atypical specimens. All this samples as well as recent collections (37 specimens) have been assimilated to this new species.
All specimens (249) deposited in BCN, LEB, MACB, MAF, PO and TSB, were examined with standard techniques by using stereoscopic and compound microscopes. A short-list was examined and photographed using a Nikon SMZ 1500 stereomicroscope and a Nikon Eclipse 80i microscope fitted with bright field (Bf) and differential interference contrast (DIC), both coupled to a Nikon DXM 1200F digital camera. To combine successive photographs at different focal levels we used the free Public Domain software CombineZ5 by Alan Hadley (http://www.hadleyweb.pwp.blueyonder.co.uk).
Secondary metabolites of all specimens were identified by thin-layer chromatography (TLC) using solvent systems A, B, C (Culberson & Kristinsson 1970, Culberson et al. 1981, Huneck & Yoshimura 1996, Orange et al. 2001). Ochrolechia parella (MACB 60317) was used as standard for the variolaric acid. Measurements (width of the branches and diameter of the soredia) were calculated by using 10 samples of each morphotype in the complex and the results are given as minimum value observed (in brackets) followed by average value obtained and maximum value observed (in brackets).
Ramalina carminae R. Arroyo & E. Serina sp. nov.
Thallo saxicolo, erecto, caespitoso vel subpendulo, rigido, usque ad 6 cm longo, cinereoflavescente. Laciniae applanatae, irregulariter divisae, ramificatione intricate, rugulosostriatae et pro parte majore dichotomae. Soraliis numerosae lateralia et terminalia, sorediis granulosi. Apotheciis ignotis. Thallo K-; medulla K+, KC+ P-; UV-. Acidi variolarici et usnici continens.
Typus: SPAIN: SORIA: Montejo de Tiermes (Archaeological Site), UTM: 30T48721 457519, 1150 m alt., N-NW exposure, sandstone, 28 August 2007, M. J. Perez Alonso (Arroyo 4284) (100.000 MACBholotypus; H, MA, UPS-isotypi) (Figs. 1-6).
Thallus shrubby, erect to subpendulous or rarely caespitose, rigid, yellow green to almost whitish in some areas which usually belong to branching points or changes in the branches width; profusely and irregularly branched. Branches with a matt and irregularly grooved surface, with veins projected specially in the lower face, irregularly shaped, frequently twisted and intermixed, giving a frayed appearance, of 1-6 cm length, (0.45-)0.78(-1.25) mm width in the base and up to (0.25-)0.58(-1.37) mm in the apices, showing a non-gradual variation and being common the expansion of the apices; cracks are common and they are frequently sorediate. Highly sorediate, especially in the upper part and particularly in the apical expansions. Soralia scattered, with a great morphological diversity, more diffuse in the apical parts, with trend toward labriform, or under the thin finger-shaped branches, more rarely capitate or helmet-shaped. Marginal and laminar soralia are better delimitated, more or less excavated. Some marginal soralia are clearly deeply concave. Others take a funnelshape covered in soredia and other capitate soralia, mainly apical, arise from thin branches by way of peduncle, almost whitish. Soredia granular, of 50-80 [micro]m diam. Pseudocyphellae sparse and oval. Apothecia andpycnidia not seen (Figs. 2-3).
[FIGURE 1 OMITTED]
Anatomy. -Like most of the species in Ramalina, it also characteristically shows a distinctly two-layered cortex. The outer cortex (paraplectenchymatous), 7-12 [micro]m thick. The inner layer chondroid cylinder (mechanical tissue prosoplechtenchymatous) forming a more or less continuous layer of uneven width, 60-250 [micro]m thick, sometimes filling the entire cross section of a branch. Medulla lax (Fig. 4).
Chemistry.-Cortex K-, C+ and KC+ weakly yellowish, P-, UV-. Medulla and soralia: K+ weakly yellowish, KC+ orange, P-, UV-. All specimens show a great chemical uniformity, containing variolaric acid as the only compound in the medulla and usnic acid in the cortex.
Etymology.-The name of this species is given in honour to Ma del Carmen Cabeza Menendez (Carmina, 19172007), mother of this work's first author. She always felt very proud of the fact that one of her daughters was dedicated to the study of something as thrilling as lichens and Ramalina genus.
Habitat and distribution .--The description of Ramalina carminae is mainly based on herborized material collected from the Tiermes archaeological site (Soria, Spain) at an altitude of 1150 m, on sandstones (type locality). These populations grow preferentially on vertical walls with N-NW exposure, protected from direct wind incidence, where R. carminae reaches a great cover; only some specimens of scarce developement were found on bird perches. In that locality Ramalina carminae grows together with the R. capitata complex, especially with R. capitata var. protecta, appearing frequently intermixed, so R. carminae may go unnoticed. Other associated species in this locality, like Ramalina farinacea (L.) Ach. and saxicolous Evernia prunastri (L.) Ach., are less common (Fig. 1). From the presence of this taxon in other Spanish localities, it can be conclude that its habitat is the saxicolous one and it grows on acid rocks in mountainous areas. The new species is most frequently found at altitude range from 1200 to 2000 m. The lowest altitude (650 m) corresponds to Beira Alta (Portugal) and the highest one (2500 m) to Penones de San Francisco, Sierra Nevada, Granada (Spain). Although there have been found some specimens over exposed siliceous rocks, Ramalina carminae is more frequent on sheltered rocky areas or on vertical walls protected from direct wind and rain. As for the distribution of Ramalina carminae, it seems to be a taxon of some mountains in the Mediterranean Region. At the present time this new species was found in several mountainous localities in Beira Alta (Portugal) and Almeria, Avila, Granada, Guadalajara, Madrid, Soria and Teruel (Spain) in the Iberian Peninsula, as well as in Sardinia (Italy) (Fig. 5).
Observations.-In the morphological range of Ramalina carminae we found two morphotypes; one of them, less usual, belongs to small rosette-shaped thallus up to 2 cm diam. and 0.5 cm tall, which can be caespitose; laciniae flat, richly dichotomously branched, distal parts fanshaped branched; these distal parts, which do not twist or do it slightly, have granular soralia on the tips. This morphotype was found on stone, in a cave of Puerto de la Morcuera (Madrid, Spain) at 1450 m. In this locality, but outside the cave, we found the first described morphotype, more common, erect to subpendulous and with a great morphological diversity of soralia, coexisting with different varieties of R. capitata and saxicolous R. pollinaria (Westr.) Ach. The small rosette-shaped morphotypes of R. carminae may recall R. capitata var. digitellata in its palmately branched finger-shaped apices of the laciniae. We studied the type material of R. digitellata Nyl. and we found differences in the branching appearance and in the curvature of the apices of the lacinae. Furthermore, R. capitata var. digitellata as well as R. polymorpha and the different varieties of R. capitata lack medullary substances, only some specimens contain norstictic acid (Arroyo et al. 1991) which causes the K+ red reaction in medulla and soralia. This reaction was also noticed by Magnusson (1956), when he described R. protecta. However R. carminae always contains variolaric acid, a very important character for the differentiation of this taxon.
[FIGURE 2 OMITTED]
The remaining samples from the other iberian localities presented in this work, were previously included into the R. capitata complex as atypical specimens with great profusion of soralia and a chemical compound with Rf (3:2:2) class as the only secondary metabolite (Arroyo et al. 1991). This material matches from a morphological, anatomical and chemical point of view with the description of this new species, that is why we assimilate all this specimens to R. carminae. Differences in the morphology of the soralia between the three varieties that we accept in the R. capitata complex are shown in Fig. 6.
[FIGURE 3 OMITTED]
Possible confusions with R. capitata var.protecta are due to the presence of some helmet-shaped soralia in R. carminae, but the rest of their morphological and chemical features are completely different. There are other probable confusion, reflected in the material of different herbaria (three specimens from Portugal, one of them stored in the Herbarium of the University of Oporto (PO) and two more stored in the Herbarium of the University of Leon (LEB 6943) Spain) that we have reviewed, between R. carminae and R. pollinaria (Westr.) Ach. and R. subfarinacea (Nyl. ex Cromb.) Nyl. Morphological as well as chemical characters of both taxons clearly differ from those of R. carminae. The specimens of R. carminae with abundant soralia may recall R. pollinaria, but in case of doubt a chemical test is decisive, because R. pollinaria always contains evernic and obtusatic acids. Confusions with R. subfarinacea are less probable despite the presence of some marginal soralia with granular soredia in R. carminae. In their morphology as well as in their chemical composition and habitat, they are very different taxa. The specimens from Sardinia (two saxicolous samples) analyzed by Dr. S. Bartsch and stored in (TSB) Herbarium Universitatis Tergestinae (Trieste, Italy) did not have any specific identification. The main differences between R. carminae and similar taxa are shown in Table 1.
Variolaric acid (orcinol depsidone) is a compound generally known in crustose lichens and frequent in Ochrolechia and Pertusaria (Zedda 1999). This compound has the peculiarity of having the ether bond in 3'position instead of 5', an exception in the biosynthesis of lichen depsidones. In the chemical analysis by TLC, a pale bluishgrey spot is observed after dilute sulphuric acid and heat in solvent A, and a pale yellowish (almost colourless) in B and C with Rf classes (3:2:2) which coincides with the unknown substance SDAM1 of Ramalina farinacea of the centre of Spain (Arroyo & Manrique 1988). In Ramalina, variolaric acid is a very infrequent compound, only reported in R. farinacea from Norway, Sweden and Spain (Mallorca Island) (T0nsberg 1982) and from Italy (Zedda 1999) and always as an accessory substance. In Ramalina, we also detected this compound in the Iberian Peninsula specimens of several chemotypes of Ramalina farinacea and in a chemotype of R. subfarinacea (Arroyo et al. unpublished). It was always occasional and an accessory substance in all cases. However we have not detected variolaric acid in the few specimens of saxicolous R. farinacea which grow close to R. carminae. The new species is, so far, the only species of the genus with variolaric acid as the only major medullary compound.
[FIGURE 4 OMITTED]
There is an important variation in the anatomical structure of R. carminae throughout the thallus (Fig. 4); some areas lack medulla and algal cells, differing only in the outside and inside cortex and they can match with a branching point or being interspersed in the laciniae; other areas show a clear predominance of the inner cortex embedded together with algal cells and medullary hyphae as isolated groups in the chondroid cylinder. There are also areas with the typical anatomical structure of Ramalina, with an inner cortex of uneven width, like a sort of "waves", algal layer more or less continuous and a loose medulla. The areas sticking out of the branches like "veins" match with a great growth of the inner cortex and a lack of algal cells. It is strange the wholly interruption of the chondroid cylinder, a frequent feature in other species of the genus Ramalina like R. fastigiata, R. fraxinea or R. polymorpha s.str. We have not observed chondroid strands embedded in the medulla, as it happens in R. capitata var.
capitata or in R. capitata var.protecta, species associated to R. carminae. The anatomy of R. carminae may recall R. tortuosa, a species which is also intricatelly branched that may superficially resemble certain eroded forms of R.capitata complex. However this species contains salazinic acid in the medulla and it is not known outside the Canary Islands (0sthagen & Krog 1976; Krog & 0sthagen 1980).
[FIGURE 5 OMITTED]
[FIGURE 6 OMITTED]
The morphological and anatomical features that characterize R. carminae clearly differ from those that characterize the different associated taxa. Besides, these differences are correlated to the presence of variolaric acid, a very infrequent compound in the genus. We consider that the specimens we present can not be treated as a chemotype of one of the varieties that we recognized in R. capitata, since this concept is related to a chemical differentiation of morphologically indistinguishable individuals of a same species (Hawksworth 1976). Based on the correlation between the morphological, chemical and anatomical differentiation, we recognize R. carminae as a new species.
From the different sites where R. carminae was collected, only in the populations of Montejo de Tiermes (Soria) there were specimens parasited with Lichenoconium usneae (Anzi) D. Hawksw., Phaeosporobolus usneae R. Sant. and Tremella ramalinae Diederich (S. Perez Ortega, per. comm.).
Additional specimens examined. ITALY: SARDINIA: 1250 m, esquistos, P.L. Nimis 1989, no 307 B&L (1992) (TSB); 1300 m, esquistos, P.L. Nimis 1989, no 304 B&L (1992) (TSb). PORTUGAL: BEIRA ALTA: 650 m, granitos, (PO 5246L). Tras-Os-Montes: 29T6861 4645, Montesinho, 1280 m, A. Terron 2006, granitos, bird perches (LEB 6943). SPAIN: ALMERIA: Bacares: 30S542 4119, Sierra de los Filabres, Merendera, 1984 m, J.M. Egea 1978, (Arroyo 1561, 1563,1564) MUB 1019 (BCN); Gergal: 30S534 4118, Sierra de los Filabres, Piedra del Sombrero, 1900 m, J.M. Egea 1982, (Arroyo 2685, 2687) (BCN). Avila: 30T319 4471, Sierra de Gredos: Navarredonda de Gredos, 1600 m, R. Arroyo & J.J. Perez 1988, (Arroyo 3237-3240, 3242, 3243, 3245-3251, 3253-3262, 3264-3266) (MACB 102148, 102150102154); 30T 3055 44593, Risco Negro, (Circo de Gredos), 2300 m, L.G. Sancho 1982, (Arroyo 809-812, 814, 816) (MACB 102147, 102149). GRANADA: Guadix: 30S531 4118, Sierra de Baza, Piedra del Deseo, 1900 m, J.M. Egea 1979, (Arroyo 1555-1559) MUB 1018 (BCN); Sierra Nevada: 30S468 4107, Penones de San Francisco, 2500 m, J.M. Egea 1980, (Arroyo 1566, 1567, 1569, 1570) MUB 1020 (BCN). GUADALAJARA: 30T59938 449553, Parque Natural Alto Tajo: Chequilla, 1360 m, E. Araujo & T. Camarero 2009, [Arroyo 4336 (13)] (MACB 102138). MADRID: 30T455 4545, Montejo de La Sierra: Hayedo de Montejo, 1300 m, R. Arroyo 1986, (Arroyo 243, 245, 247251) (MACB 102156, 102157). 30T407 4511, Sierra de Guadarrama: La Penota, 1600 m, L.G. Sancho 1978, (Arroyo 765-772) (MACB 102158); 30T402 4497, El Escorial, Monte Abantos, 1754 m, R. Arroyo & J. J. Perez 1988, (Arroyo 2491-2530, 2532-2544) (MACB 102140102145); 30T431 4519, Puerto de la Morcuera, 1450 m, A. Pintado 2006, s/n (MAF); A. Pintado & R. Arroyo 2006 [Arroyo 4327, 4328, 4330, 4331 (1,2), 4332(1-5)] (MACB 102132-102136). 30T42 451, Manzanares El Real, La Pedriza de Manzanares (al pie de Cabezas de Hierro), 1900 m, granitos protegidos, E. Manrique 1986, (Arroyo 699704) (MACB 102146). SORIA: 30TVL8479, Montejo de Tiermes: pared vertical, arcillas rojas expuestas, 1200 m, I. Perez 1987, (Arroyo 1828,1830) (MACB 102155); 30T48721 4575195, Yacimiento Arqueologico, areniscas protegidas, 1150 m, M. J. Perez 2007, [(Arroyo 4221 (3-11)] (MACB 102121); R. Arroyo, E. Serina, & E. Araujo 2008, [Arroyo 4266'(2), 4267'(1-6), 4268'(1-4), 4269'(1-6), 4270'(1-8), 4271'(27), 4272'(1-7), 4273(1-7), 4274(1-4), 4275(1-4), 4276(5)] (MACB 102122-102131, 102139); E. Araujo & M. Cervera 2009, [Arroyo 4335(1-9)] (MACB 102137). TERUEL: Sierra de Albarracin: Orihuela del Tremedal, 1500 m, X. Llimona 1986, (Arroyo 1927-1936) (BCN 625); 30TXK159808, Noguera de Albarracin hacia Orihuela del Tremedal, area recreativa Pena del Castillo, piton diacitico, 1580 m, R. Arroyo, E. Serina & E. Araujo 2010 (MACB 102058-102064); 30TXK143874, Orihuela del Tremedal, Santuario Ntra. Sra. del Tremedal, cuarcitas, 1725 m, R. Arroyo, E. Serina & E. Araujo 2010 (MACB 102066-102070); 30TXK197845, Bronchales, hacia Noguera, cuarcitas, 1606 m, R. Arroyo, E. Serina & E. Araujo 2010 (MACB 102076).
The authors are grateful to Ma Jose Perez Alonso for collecting the first specimens in Montejo de Tiermes (type locality). Victor J. Rico is thanked for helping with the photographic subjet and for his valuable comments. We would like to express our thanks to P. L Nimis and Elisa Folhadela for providing the possibility of checking the material in the Herbaria (TSB) Herbarium Universitatis Tergestinae (Trieste, Italy) and (PO) University of Oporto. Blanca Gonzalez helped with her advices in staining microscopy. Pradeep K. Divakar is gratefully acknowledged for revising the English text. This research was supported by a grant from the Spanish DGICYT (CGL2004-04795-C0401/BOS and CGL2007-66734-C03-01/BOS).
Recibido: 26 mayo 2011. Aceptado: 21 junio 2011
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Rosario Arroyo, Estela Serina and Elena Araujo (1)
(1) Departamento de Biologia Vegetal I, Facultad de Biologia, Universidad Complutense, E-28040 Madrid, Spain. firstname.lastname@example.org; email@example.com
Tabla 1 Main differences between Ramalina carminae and similar taxa Character R. carminae Branches width: Basal (0.45-) 0.78 (-1.25 mm) Middle (0.42-) 0.85 (-1.20) mm Apical (0.25-) 0.58 (-1.37) mm Branching Irregularly branched, frequently twisted Soralia distribution Scattered throughout the thallus Soralia morphology Great morphological diversity Soredia 50-80 urn Apothecia Not seen Anatomy Variation throughout the thallus Chondroid strands embedded in the medulla not seen Chemistry Variolaric and usnic acids Habitat Sheltered acid rocks 1200-2000 (2500) m Character R. capitata var. capitata Branches width: (087-) 1.78 (-3.62) mm (0.75-) 1.65 (-3.75) mm (0.70-) 1.008 (-2.37) mm Branching Regularly and palmately branched Soralia distribution Apical Soralia morphology Predominantly capitate or subcapitate, some labriform Soredia 80-120 [micro]m Apothecia Rare, terminal- subterminal Anatomy Continuous inside cortex of uneven width Chondroid strands embedded in the medulla Chemistry Usnic acid (exceptionally norstictic acid) Habitat Exposed acid rocks (100-) 1000-15000 (-2500) m Character R. capitata var. protecta Branches width: (1.00-) 2.05 (-3.50) mm (0.75-) 2.25 (-3.50) mm (0.50-) 1.50 (-2.62) mm Branching Regularly and pallmately branched Soralia distribution Apical Soralia morphology Predominantly helmet- shaped Soredia 40-60 [micro]m Apothecia Rare, terminal- subterminal Anatomy Continuos inside cortex of uneven width Chondroid strands embedded in the medulla Chemistry Usnic acid (exceptionally norstictic acid) Habitat Predominantly on sheltered acid rocks (100-) 1000-15000 (-2500) m Character R. capitata var. digitellata Branches width: (0.87-) 1.30 (-1.75) mm (1.12-) 1.46 (-1.75) mm (0.50-) 0.85 (-1.12) mm Branching Regularly and palmately branched Soralia distribution Apical Soralia morphology Under thin finger-shaped branches Soredia 50-60 [micro]m Apothecia Not seen Anatomy Discontinuous inside cortex of uneven width Chondroid strands embedded in the medulla not seen Chemistry Usnic acid (exceptionally norstictic acid) Habitat Predominantly on sheltered acid rocks (100-) 1000- 15000 (-2500) m
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|Author:||Arroyo, Rosario; Serina, Estela; Araujo, Elena|
|Date:||Jan 1, 2011|
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