SciELO - Scientific Electronic Library Online

 
vol.43 número3-4Hongos micorrícico arbusculares presentes en bosques de Alnus acuminata (Betulaceae) de la Yunga ArgentinaEspectro de hospedadores hallado en Beauveria bassiana y Metarhizium anisopliae (Ascomycota: Hypocreales) en Argentina índice de autoresíndice de materiabúsqueda de artículos
Home Pagelista alfabética de revistas  

Servicios Personalizados

Revista

Articulo

Indicadores

  • No hay articulos citadosCitado por SciELO

Links relacionados

Compartir


Boletín de la Sociedad Argentina de Botánica

versión On-line ISSN 1851-2372

Bol. Soc. Argent. Bot. v.43 n.3-4 Córdoba ago./dic. 2008

 

New records in the lichen family Verrucariaceae (Ascomycota) from Argentina

 

M. Prieto1 *, G. Aragón1 y I. Martínez1

1Área de Biodiversidad y Conservación, Departamento de Biología y Geología, Escuela Superior de Ciencias Experimentales y Tecnología, Universidad Rey Juan Carlos, c/ Tulipán s/n, 28933 Móstoles, Madrid (Spain).
*e-mail: maria.prieto@urjc.es.

 


Summary: Four interesting lichens belonging to Catapyrenium, Placidium and Placopyrenium genera (Verrucariaceae) were collected for the first time in Argentina. Catapyrenium exaratum was only reported from Chile and Perú. Placidium acarosporoides was previously known from North America, Chile and South Africa. Placidium pilosellum, a widespread and common species, but little known in South America. Finally, Placopyrenium bucekii, a mediterranean-submediterranean lichen, represents a new record from South America. For these taxa we provide a brief description with emphasis on the ecological aspects and distribution maps.

Key words: Lichens; Verrucariaceae; Argentina; New records.

Resumen: Nuevos registros en la familia de líquenes Verrucariaceae (Ascomycota) de Argentina. En el presente trabajo se dan a conocer cuatro especies liquénicas que constituyen novedades para Argentina. Catapyrenium exaratum, previamente recolectada en Chile y Perú. Placidium acarosporoides, conocida hasta el momento de Norte América, Chile y Sudáfrica. Placidium pilosellum, una especie común y ampliamente distribuida, pero poco conocida en Sudamérica. Finalmente, Placopyrenium bucekii, un liquen (sub)-mediterráneo ha sido recolectado por primera vez en Sudamérica. Se añaden, para cada especie, una breve descripción morfo-anatómica y ecológica y un mapa de distribución geográfica.

Palabras clave: Líquenes; Verrucariaceae; Argentina; Nuevas citas.


 

Introducción

In this paper we present four new records of pyrenocarpous lichens collected in Argentina during 2005. Three of them are new records from Argentina and the other is collected for the first time in South America. All of them are members of the lichen family Verrucariaceae, characterized by perithecial ascomata, bitunicate asci, hamathecium lacking paraphyses (at least at the maturity) but formed by periphyses and pseudoparaphyses (Janex-Favre, 1971), and a positive reaction of the hymenial gel to potassium-iodine (Henssen & Jahns, 1974).
The recently published checklist of Argentinian lichens (Calvelo & Liberatore, 2002) in which 1670 species were recorded, provided a basis for taxonomic
work on lichens in the area. This work constitutes an update of the information given by Grassi in 1950 by adding the information contained in posterior reviews as well as in different taxonomical revisions.
Very few works have been published on Verrucariaceae in South America. Although some papers have reported new records or described new taxa on different members of the family (e. g.: Räsanen, 1938, 1939; Lamb, 1955, 1958; Osorio & Ferraro, 1975; Scutari et al., 2002; Prieto et al., 2007), till the moment there hasn´t been done any treatment about the family. A first approximation was done by Breuss (1993a) who reported 13 species of Catapyrenium s.l. from South America, 10 of them from Argentina. Later, the same author, in 1995, presented a preliminary survey of Catapyrenium species occurring in the Southern Hemisphere with special emphasis on the distribution patterns. The present study aims at contributing to the knowledge of the ecology and distribution of these rare species in Argentina, although a deeper work is necessary.

Materials and Methods

The study is based on fresh material collected by the authors and included in MA and BCRU herbaria. Several specimens were requested from other herbaria (ABL and LI) and from personal collection of the authors in order to check the samples. Distributional maps were drawn using a UTM grid (10 x 10 km) projection with Arcview GIS v. 3.1.

Results

Catapyrenium exaratum Breuss, Plant Syst. Evol. 185: 26 (1993)
Illustration : Breuss 1993a, p. 27.
Specimens examined. ARGENTINA: Prov. Salta, P. N. Los Cardones, 3700 m, on calcareous soils, 01/ 08/2005, G. Aragón & I. Martínez (BCRU).
Other specimens examined. ECUADOR: Prov. Cotopaxi, P. N. Cotopaxi, along road from Pampa de Limpios to summit of Cotopaxi, 00º38´S, 78º26´W, 4100 m, grass paramo with scattered shrubs, 17/12/ 1983, R.C. Harris (LI). Typus: PERÚ: Highland from Cerro de Pasco-Carpish-Tingo Maria, 9/11/1960, F. Mattick (LI).
Other specimens examined: C. cinereum. SPAIN: Asturias, Teverga, Puerto de Ventana, Huerto del Diablo, 256962 N, 4772777 E, 1872 m, on limestone soils in alpine grassland vegetation, 18/05/2006, M. Prieto (MA). Cantabria, P. N. Picos de Europa, Orcados Rojos, 351425 N, 4782493 E, 2350 m, outcrops of compact sandstones with cuarcites 05/ 10/2005, G. Aragón, A. García & M. Prieto (MA). Huesca, Cerler, ski resort, slope Tubo de la Cogulla, 790430 N, 4721834 E, 2300 m, on soils in alpine grassland vegetation, 01/04/2006, M. Prieto (MA).

Thallus of scattered to adjacent squamules, not overlapping, to 6 mm wide, rounded to slightly lobed, totally adpressed to the substrate; upper surface pale, matte, covered by a whitish to gray pruina. Perithecia immersed and frequent. Pycnidia not found. Thallus 200-300 μm thick; upper cortex thin and poorly delimited from the algas layer, with roundish angular small cells (called cinereum-type by Breuss, 1990); epinecral layer 25-40 μm thick, cracked into subconical portions thus giving the surface an areolate appearance; algal layer 187-212 μm thick, algal cells 3-12 μm diam.; medulla lacking. Lower cortex well differentiated, 25-40 μm thick, of roundish-angular cells 4-7 μm diam. Rhizohyphae hyaline to brownish, 2.5-4 μm thick, densely interwoven giving an aspect of a rhizine. Perithecia laminally immersed, appearing as black dots on upper surface, subglobose, exciple pale. Ascospores simple, colorless, biseriately arranged, 8 per ascus, 12-20 × 7-9 μm.
The species is similar to C. cinereum (Koerber) Pers., from which it differs in having a roughly cracked necral layer, pale exciple and smaller spores. Also it could be confused with C. squamellum (Nyl.) Thomson, which has bundles of dark rizohyphae and a differentiated medullary tissue while C. exaratum develops faintly brown pigmented rizohyphae and lacks medulla.
This species grows at high altitudes (3700 to 4100 m alt), on calcareous soils together with Placidium andicola (Breuss) Breuss.
Catapyrenium exaratum has been previously reported from Ecuador and Perú (Breuss 1993a). This record extends the distribution area of the species in South America confirming the Andean distribution of the species (Fig. 1).


Fig. 1. Distribution of Catapyrenium exaratum. new records, previous records.

Placidium acarosporoides (Zahlbr.) Breuss, Bull. Cal. Lich. Soc. 7: 39 (2000)
Illustration: Thomson 1987, p. 29.
Specimens examined. ARGENTINA: Prov. Salta, Iruya, 3800 m, on limestone soils, 7/08/2005, G. Aragón & I. Martínez (MA).
Other specimens examined: MEXICO: Baja California, 1 km N of Cataviña, northen Vizcaíno Region of the Sonoran Desert, 29º43´N/114º40´W, 560 m, batholitic granite boulder area, stream with Washingtonia, (ABL). USA: Arizona: Yuma Co, Palm Canyon, 1000 ft, on rock, 1973, T. H. Nash (LI 297542). Mohave: 20 miles of Kigman, Union Pass, Black mts along Rte 68, 3600 ft, on schist, 1973, T. H. Nash (LI).

Thallus of convex contiguous squamules, up to 2 mm diam., rounded to slightly lobed; upper surface brown to red-brown, smooth. Thallus 230-600 μm thick; upper cortex 25-60 μm thick, paraplectenchymatous; epinecral layer up to 20 μm thick; algal layer 70-150 μm thick, algal cells 12-15 μm in diam.; medulla prosoplectenchymatous. Lower cortex weakly differentiated, with more densely aggregated cells; rhizohyphae hyaline 3-6 μm thick, confined to the central area, giving an aspect of a peduncle. Perithecia laminally immersed, few per squamule, subglobose, exciple colorless to brown, asci cylindrical to clavate. Ascospores simple, broadly ellipsoidal, colorless, (sub)-biseriately, 8 per ascus, 12-17 × 7-11 μm. Pycnidia laminally immersed, Dermatocarpon type, conidia oblong to shortly cylindrical.
This is a characteristic species with convex to almost bullate squamules that forms an areolate-appearing thallus.
It has been found growing on calcareous soils in Festuca grassland associated with P. andicola and P. squamulosum var. argentinum (Räsänen) Breuss.
Placidium acarosporoides was previously reported from granitic rocks and sandstones from the sea level up to 1890 metres altitude. However, in Argentina this taxon grows on calcareous soils at 3800 metres altitude. This species seems to have a disjunct distribution with occurrences in SW North America (Thomson 1987, 1989), Argentina (this paper), Chile (Breuss 1993a) and South Africa (Breuss 1993b) (Fig. 2).


Fig. 2. Distribution of Placidium acarosporoides. new records, previous records.

Placidium pilosellum (Breuss) Breuss, Annln. naturh. Mus. Wien, Ser. B, Bot. Zool. 98(Suppl.): 39 (1996)
Illustration: Breuss 1990, fig. 18.
Specimens examined. ARGENTINA: Prov. Jujuy, Iruya, Quebrada de Humahuaca, 4100 m, on limestone soils, 07/08/2005, G. Aragón & I. Martínez (BCRU).
Other specimens examined : MEXICO : Baja California, 10 km. N el Rosario in canon del Rosario, 90 m, coastal scrub community, 05/01/1989, J. M. Egea (LI).

Thallus of adjacent to overlapping squamules, to 6 mm diam., rounded to lobed, with the margins free from the substrate; upper surface orange brown, tan or dark brown; lower surface usually pale, sometimes brown to black. Perithecia immersed and frequent. Pycnidia marginal. Thallus thin, 250-400 μm thick; upper cortex 40-80 μm thick, paraplectenchymatous; epinecral layer up to 50 μm thick; algal layer 80-140 μm thick; medulla with many spherical cells, with cells of 9-12 μm diam. Lower cortex weakly differentiated, with more densely aggregated cells; rhizohyphae hyaline 4.5-6 μm thick. Perithecia laminally immersed, pyriform, exciple colorless, asci cylindrical. Ascospores simple, ellipsoidal, colorless, uniseriately arranged, 8 per ascus, 12-17 × 5.5-7.5 μm. Pycnidia marginal, Dermatocarpon type, conidia 3-4 × 1-1.8 μm.
This species is characterized by its marginal pycnidia (character that separates this species from P. squamulosum (Ach.) Breuss) and a medulla with many globular cells. Anatomically, P. pilosellum is very similar to P. andicola. Although the discrimination is very difficult, P. pilosellum has larger spores (12-17 x 5.5-7.5 μm versus 11-14 x 5-6.5 μm) and squamules with free margins meanwhile P. andicola has more or less dark rimmed squamules (Breuss 1993a).
We found it growing on limestone soils, dominated by grassland vegetation with Stipa spp. and Festuca spp. It was found together with P. squamulosum var. argentinum and Anthracocarpon andinum Prieto, Aragón & Breuss.
Placidium pilosellum is a widespread and very common species in Europe (e. g.: Breuss 1990, 1998, Grube et al. 2001, Hafellner & Türk 2001, Llimona& Hladun 2001). It has been also found in Asia (Breuss 1998, Seaward et al. 2004), Australia (Breuss 2001), North America (Nash et al. 1998), and South America (Feuerer & Sipman 2005). This is the second record
of the species in South America, being previously known from Bolivia.

Placopyrenium bucekii (Nádv. & Servít) Breuss, Stud. Geobot. 7: 182 (1987)
Illustration: Breuss 1993c p. 9; Ménard & Roux 1995.
Specimens examined. ARGENTINA: Prov. Salta, P. N. Los Cardones, 3700 m, on fissures of calcareous rocks, 01/08/2005, G. Aragón & I. Martínez (MA).
Other specimens examined: P. bucekii var. bucekii. TURKEY: Bursa, 2003, on rock, (ABL). SPAIN: Canary Inslands, Tenerife, Macizo Anaga, slope near Semáforo de San Andrés, 80-150 m., 10/ 07/1986, O. Breuss (LI). P. bucekii var. triseptatum. TURKEY: Prov. Izmir, Yamanlardað, southern flank above Yamanlarköy, along the path to Karagöl, 38º32´N/27º09´E, 700 m, 14/04/1992, O. Breuss (LI).

Thallus epilithic, squamulose-areoled, the squamules are morphologically different according to the position in the thallus, being polygonal the central ones and lobed the most peripherical squamules. All of them are covered by a whitish to grey pruine. Perithecia are immersed in the squamules and are detectable only by their top, forming a black point. Studied material belongs to var. triseptatum Breuss. This variety is recognized by its 3-celled spores, whereas the variety type has 1- or 2-celled
spores. Spore size is 16-23 x 6-8 μm in var. triseptatum (versus 13.5-17 x 5-7 μm in var. type). Placopyrenium bucekii has a mediterranean-submediterranean distribution with occurrences in the Canary Islands and Caucasus (Breuss 1987). It has been also cited in Albania (Hafellner & Kashta 2003), Cyprus, Greece and France (Ménard & Roux 1995), Iran (Seaward et
al. 2004), Israel (Galun & Mukhtar 1996) and Turkey (Breuss 1993c, John 1996). Previously var. triseptatum was only known from Turkey, growing on calcareous soils at 700 m altitude (Breuss 1993c). We have found it in Argentina on fissures of rocks together with C. exaratum and P. andicola.
The new record extends its distribution range to South America and increases its altitudinal range to more than 3000 m (Fig. 3). So far, this variety might have gone unnoticed and probably it will reach a wider distribution.


Fig. 3. Distribution of Placopyrenium bucekii. new records, previous records.

Addenda

We include here another species collected in Argentina although they are not new records for the country in order to complete the knowledge about the distributional patterns.

Placidium andicola (Breuss) Breuss, Annln. naturh. Mus. Wien, Ser. B, Bot. Zool. 98(Suppl.): 38 (1996)
Illustration: Breuss 1993a, p. 21.
ARGENTINA: Prov. Jujuy: Hornaditas, Quebrada de Humahuaca, 3260 m, puna vegetation, 13/02/2007, A.R. Burgaz. Prov. Salta, Iruya, 3800 m, on limestone soils, 7/08/2005, G. Aragón & I. Martínez (MA). P. N. Los Cardones, 3700 m, on calcareous soils, 01/08/2005, G. Aragón & I. Martínez (MA).
Other specimens examined : MEXICO : Baja California, 10 km. N el Rosario in canon del Rosario, 90 m, coastal scrub community, 05/01/1989, J. M. Egea (LI).

Thallus of dispersed to adjacent squamules, to 4 mm diam., rounded to lobed, with the margins slightly ascending; upper surface brown, black-rimmed; lower surface usually dark brown to black. Perithecia immersed and frequent. Pycnidia marginal. Thallus 260-400 μm thick; upper cortex 35-75 μm thick, paraplectenchymatous; epinecral layer up to 15 μm thick; algal layer 75-120 μm thick, algal cells 3-9 μm in diam.; medulla with many spherical cells. Lower cortex weakly differentiated, with more densely aggregated cells; rhizohyphae hyaline to brown 4.5-6.5 μm thick. Perithecia laminally immersed, exciple colorless, darkening with the age, asci cylindrical. Ascospores simple, ellipsoidal, colorless, uniseriately arranged, 8 per ascus, 10-14 × 5-6.5 μm. Pycnidia marginal, Dermatocarpon type, conidia 2.5-4 × 1-1.8 μm.
This is a relatively common species in South America, previously reported from Argentina, Bolivia and Peru (Breuss 1993a). We have found it at high altitude on calcareous soils together with C. exaratum, P. acarosporoides, P. squamulosum var. argetinum and P. bucekii var. triseptatum.

Placidium squamulosum var. argentinum (Räsänen) Breuss, Annln. naturh. Mus. Wien, Ser. B, Bot. Zool. 98(Suppl.): 39 (1996)
Illustration: Breuss 1990 fig. 20.
ARGENTINA: Prov. Salta, Salina Grande, on limestone soils, 3310 m, 5/08/2005, G. Aragón & I. Martínez (MA). Prov. Salta, Iruya, on limestone soils, 3800 m, 7/08/2005, G. Aragón & I. Martínez (MA).

Thallus of adjacent to slightly overlapping squamules, to 7 mm diam., rounded to lobed; upper surface brown; lower surface pale to brown. Perithecia immersed and frequent. Pycnidia laminal. Thallus 200- 400 μm thick; upper cortex 30-80 μm thick, paraplectenchymatous; epinecral layer up to 50 μm thick; medulla with many spherical cells, of 9-14 μm diam. Lower cortex weakly differentiated, with more densely aggregated cells; rhizohyphae hyaline 4-5 μm thick. Perithecia laminally immersed, pyriform, exciple colorless, asci cylindrical. Ascospores simple, ellipsoidal, colorless, uniseriately arranged, 8 per ascus, 14-16 × 7-9 μm. Pycnidia laminal, Dermatocarpon type, conidia oblong-ellipsoid.
This variety differs from the type one in having broader spores and thinner rizohyphae (spores 12-16 × 5.5-7.5 μm and rizohyphae 4.5-6.5 μm in var. squamulosum). It has a very restricted distribution, known from Argentina (Breuss 1993a) and few collections from Central and North America (unpub. Breuss). We have found it in Argentina growing on calcareous soils in Festuca grassland associated with P. andicola and P. acarosporoides.

Acknowledgments

We express our sincere thanks to the curators of the herbaria ABL and LI who sent us the material for the study. We are also grateful to O. Breuss for his invaluable help. This study was supported by the Spanish «Ministerio de Educación y Ciencia» (project CGL2004-04795-C04-04) and by a PhD. grant (Universidad Rey Juan Carlos) to M. Prieto.

Bibliography

1. BREUSS, O. 1987. Placopyrenium gen. nov. In: Nimis, P.L., Poelt, J. (eds) The lichens and lichenicolous fungi of Sardinia (Italy). Stud. Geobot. 7: 182-183.         [ Links ]

2. BREUSS, O. 1990. Die Flechtengattung Catapyrenium (Verrucariaceae) in Europa. Stapfia 23: 1-153.         [ Links ]

3. BREUSS, O. 1993a. Catapyrenium (Verrucariaceae) species from South America. Plant Syst. Evol. 185: 17-33.         [ Links ]

4. BREUSS, O. 1993b. Studien über die Flechtengattung Catapyrenium (Verrucariaceae). V. Einige Arten aus dem südlichen Afrika. Linzer biol. Beitr. 25: 339-346.         [ Links ]

5. BREUSS, O. 1993c. Zwei neue Flechtentaxa aus der Türkei. Österr. Z. Pilzk. 2: 7-10.         [ Links ]

6. BREUSS, O. 1995. The genus Catapyrenium (Verrucariales) in the southern hemisphere. Crypt. Bot. 5: 177-183.         [ Links ]

7. BREUSS, O. 1998. Catapyrenium und verwandte Gattungen (lichenisierte Ascomyceten, Verrucariaceae) in Asien - ein erster Überblick. Annaln. Naturh. Mus. Wien 100B: 657-669.         [ Links ]

8. BREUSS, O. 2001. Placidium. In: McCarthy, P.M. (ed.): Flora of Australia. Vol. 58 A, Lichens 3: 168-170. ABRS/CSIRO Australia, Melbourne.         [ Links ]

9. CALVELO, S. & S. LIBERATORE. 2002. Catálogo de los líquenes de la Argentina. Kurtziana 29: 7-170.         [ Links ]

10. FEUERER, T. & H. J. M. SIPMAN. 2005. Additions to the lichenized and lichenicolous fungi of Bolivia. Herzogia 18: 139-144.         [ Links ]

11. GALUN, M. & A. MUKHTAR. 1996. Checklist of the lichens of Israel. Bocconea 6: 149-171.         [ Links ]

12. GRASSI, M. M. 1950. Contribución al catálogo de líquenes argentinos, I. Lilloa 24: 5-296.         [ Links ]

13. GRUBE, M., L. LINDBLOM & H. MAYRHOFER. 2001. Contributions to the lichen flora of Crete: A compilation of references and some new records. Stud. Geobot. 20: 41-59.         [ Links ]

14. HAFELLNER, J. & R. TÜRK. 2001. Die lichenisierten Pilze Österreichs - eine Checkliste der bisher nachgewiesenen Arten mit Verbreitungsangaben. Stapfia 76: 1-167.         [ Links ]

15. HAFELLNER, J. & L. KASHTA. 2003. Miscellaneous records of lichens and lichenicolous fungi from Albania. Herzogia 16: 135-142.         [ Links ]

16. HENSSEN, A. & H. M. JAHNS. 1974. Lichenes. Eine Einführung in die Flechtenkunde. Georg Thieme Verlag, Stuttgart.         [ Links ]

17. JANEX-FAVRE, M. C. 1971. Recherches sur l'ontogénie, l'organisation et les asques de quelques pyrénolichens. Rev. Bryol. Lichénol. 37: 421-469.         [ Links ]

18. JOHN, V. 1996. Preliminary catalogue of lichenized and lichenicolous fungi of Mediterranean Turkey. Bocconea 6: 173-216.         [ Links ]

19. LAMB, I. M. 1955. New lichens from northern Patagonia, with notes on some related species. Farlowia 4: 423- 471.         [ Links ]

20. LAMB, I. M. 1958. La vegetacion liquénica de los Parques Nacional Patagónicos. An. de Parques Nacionales 7: 1-188.         [ Links ]

21. LLIMONA, X. & N. L. HLADUN. 2001. Checklist of the lichens and lichenicolous fungi of the Iberian Peninsula and Balearic Islands. Bocconea 14: 1-581.         [ Links ]

22. MÉNARD, T. & C. ROUX. 1995. Placopyrenium bucekii et remarques sur les Verrucariaceae (Verrucariales, Lichenes). Mycotaxon 53: 129-159.         [ Links ]

23. NASH III, T.H, B. D. RYAN, W. C. DAVIS, O. BREUSS, J. HAFELLNER, H. T. LUMBSCH, L. TIBELL & T. FEUERER. 1998. Additions to the lichen flora of Arizona IV. Bryologist 101: 93-99.         [ Links ]

24. OSORIO, H. S. & L. I. FERRARO. 1975. Contribution to the lichen flora of Argentina. VII. New and noteworthy records from the province of Corrientes. Comun. Bot. Mus. Hist. Nat. Montevideo 57: 1-4.         [ Links ]

25. PRIETO M., G. ARAGÓN, I. MARTÍNEZ & O. BREUSS. 2008. A new species of the lichen genus Anthracocarpon (Verrucariaceae) from Argentina. Bryologist 111: 128-132.         [ Links ]

26. RÄSÄNEN, V. 1938. Beiträge zur Flechtenflora Südamerikas. Rev. Sudamer. Bot. 5: 65-72.         [ Links ]

27. RÄSÄNEN, V. 1939. Contribución a la flora liquenológica sudamericana. An. Soc. Sci. Arg. 128: 133-147.         [ Links ]

28. SCUTARI, NC, B. D. DIEZ & H. T. LUMBSCH. 2002. New records of soil-associated lichens from north-eastern Patagonia (Chubut, Argentina). Mycotaxon 83: 369-383.         [ Links ]

29. SEAWARD, M. R. D., H. J. M. SIPMAN, M. SCHULTZ, A. A. MAASSOUMI, M. H. M. ANBARAN & M. SOHRABI. 2004. A preliminary lichen checklist for Iran. Wildenowia 34: 543-576.         [ Links ]

30. THOMSON, J. W. 1987. The lichen Genera Catapyrenium and Placidiopsis in North America. Bryologist 90: 27-39.         [ Links ]

31. THOMSON, J. W. 1989. Additions and a Revised Key to Catapyrenium in North America. Bryologist 92: 190-193.         [ Links ]

Recibido el 03 de Enero del 2008,
aceptado el 13 de junio, del 2008.

Creative Commons License Todo el contenido de esta revista, excepto dónde está identificado, está bajo una Licencia Creative Commons