Introduction
While revising the species of the genus Rinodina (Ach.) Gray belonging to the Dolichospora group (at present including R. brasiliensis Giralt, Kalb & H. Mayrhofer, R. dolichospora Malme, R. guianensis Aptroot, R. intermedia Bagl. and R. inspersoparietata Giralt & van den Boom), typically characterized by containing drops of uncertain origin and nature surrounding the lumina of the ascospores (Giralt et al. Reference Giralt and van den Boom2008, Reference Fos and Giralt2009), we examined several muscicolous, isidiate Rinodina specimens from Venezuela collected at high altitude in open Paramo vegetation. These specimens were somewhat similar in habit to the isidiate but corticolous R. brasiliensis occurring in tropical rainforests, but, because of their different ecological behaviour we suspected that they belonged to a new Rinodina species. Detailed morphological and chemical studies have shown that they can be clearly distinguished from R. brasiliensis by the absence of inclusions in the ascospores and by an alternative chemistry.
Materials and Methods
The specimens were examined by standard techniques using stereoscopic and compound microscopes. Current mycological terminology generally follows Kirk et al. (Reference Kirk, Cannon, David and Stalpers2001). Only free ascospores lying outside the asci have been measured. Measurements were made in water at ×1000 magnification. Mean value (x) and standard deviation (SD) were calculated and the results are given as (minimum value observed) x ± SD (maximum value observed) followed by x, SD and n (the total number of ascospores measured) in parentheses. The terminology used for the asci follows Rambold et al. (Reference Rambold, Mayrhofer and Matzer1994) and for the ascospore-types and ascospore-ontogenies Giralt (Reference Giralt2001). Chemical constituents were identified by thin-layer chromatography (TLC) and high performance liquid chromatography (HPLC) (Elix et al. Reference Elix, Giralt and Wardlaw2003).
The Species
Rinodina fuscoisidiata Giralt, Kalb & Elix sp. nov
Rinodinae guianensis similis, sed thallo et terpenum ignotum et atranorinum continenti, isidiis majoribus, c. 0·07–0·2 × 0·07–1 mm, apotheciis lecanorinis et majoribus, c. (0·5–)0·7–0·9(–1·2) mm latis, et ascosporis majoribus, c. (17–)20–25(–29) × (9–)10·3–12·7(–14) μm differt.
Typus: Venezuela, Mérida, distr. Rangel, zwischen Laguna Mucubaji und Pico Mucuñuque, etwa 15 km SE von Apartaderos. In Paramo-Vegetation, 3500 m, 8°45′N; 70°45′W, 7, 8 & 16 August 1989, K. & A. Kalb & López-Figueiras (hb. Kalb 27013—holotypus; GZU, hb. Kalb 27013—isotypus).
Thallus muscicolous, crustose, densely isidiate, spreading. Isidia simple and globose at first, 0·07–0·2 mm diam., becoming cylindrical, simple or more commonly branched or coralloid, up to 1 mm long, pale chestnut brown to dark brown, darker and often erumpent at the apices, not forming soredia or blastidia (Fig. 1). Cortex paraplectenchymatous, 20–35 μm thick, composed of mesodermatous hyphae with rounded to ±elongate cells 7–10 μm (textura globularis to angularis), chestnut brown pigmented in the outermost part, colourless in the inner part, totally interspersed with crystals soluble in K. Medulla 60–130 μm, I+ pale blue, totally interspersed with crystals soluble in K. Algal cells chlorococcoid, 8–15 μm diam.

Fig. 1: Rinodina fuscoisidiata, habitus showing the isidiate thallus and the apothecia with plane discs and prominent thalline margins (holotype). Scale = 1mm.
Apothecia lecanorine, sessile to shortly stipitate, markedly constricted at the base, with a short brown stipe, usually scattered, (0·5–)0·7–0·9(–1·2) mm diam. Thalline margin concolorous with isidia, thick, prominent, entire to verrucose, becoming isidate, sometimes partially excluded. Disc dark brown, concave to flat, funnel-shaped with age, epruinose. Proper margin entire, thin, clearly visible when the thalline margin is partially excluded. Thalline exciple 80–120(–140) μm wide, totally interspersed with crystals soluble in K; cortex paraplectenchymatous, 10–20 μm wide in the lateral part, expanded to 40–50(–80) μm below, I+ pale blue. Proper exciple indistinct laterally to 25 μm thick, expanded to 35–50(–70) μm above. Hymenium colourless, 100–120 μm high; epihymenium chestnut brown. Hypothecium colourless, 90–125 μm deep. Paraphyses 1–1·5 μm wide, apical cells 3–5 μm wide, with brown cap. Asci Lecanora-type, 8-spored. Ascospores Pachysporaria-type, (17–)20–25(–29) × (9–)10·3–12·7(–14) μm (M= 22·5 × 11·5 μm; SD= 2·5/1·2 μm; n= 75), with irregularly rounded lumina, sometimes with protrusions towards the spore ends, when mature slightly constricted at septum, walls not ornamented, torus absent or poorly developed, ontogeny of type B; many simple, colourless ascospores with apical wall thickenings are always present; when overmature a few ascospores show an additional apical lumina in each cell (Fig. 2).

Fig. 2. Rinodina fuscoisidiata, ascospore ontogeny type B and ascospore variability (holotype). Scale = 10 μm.
Pycnidia and conidia not seen.
Chemistry. Thallus, isidia and thalline margin K+ yellow, C−, KC−, Pd−, UV−. Containing atranorin [minor] and an unknown terpene [major] by TLC, HPLC. Because of the brown colour of the thalline cortex the K+ yellow reaction can be seen more easily under the microscope. After the application of K, the crystals present in the cortex and the medulla dissolve giving a yellowish coloration; after a few seconds, colourless, acicular crystals 4·5-10 μm long develop from the lichen tissue and these later regroup to form star-like structures. When K was applied to the moss substratum no crystals developed.
Ecology and distribution. The new species is known only from Merida province in Venezuela. It is growing over mosses on humid granitic boulders in open paramo vegetation between 3500 and 4200 m. Accompanying species are Tetramelas regiomontanus Marbach, Rinodina stictica Sheard & Tønsberg and Buellia aff. proximata.
Observations. Rinodina fuscoisidiata is characterized by its continuous, brown, densely isidiate thallus, the presence of an unknown terpene (major) and atranorin (minor), the thick, long, simple or ramified to coralloid isidia and by the large Pachysporaria-type ascospores. The ascospores are smooth, slightly constricted at the septum, with a poorly developed torus or torus absent and develop with type B ontogeny.
There are only four other truly isidiate species of Rinodina: the corticolous/muscicolous R. brasiliensis Giralt, Kalb & H. Mayrhofer, R. guianensis Aptroot, R. isidioides (Borrer) H. Olivier and the saxicolous R. placynthielloides Aptroot. The tropical to subtropical R. brasiliensis and R. guianensis also have brownish isidia and Pachysporaria-type ascospores but lack secondary lichen substances. Furthermore their ascospores contain minute globular inclusions surrounding the lumina (compare Giralt et al. Reference Giralt, Kalb and Mayrhofer2009). The oceanic-temperate R. isidioides has whitish to pale grey isidia united at the base into a subsquamulose structure and contains only atranorin (Sheard Reference Sheard1967; Giralt et al. Reference Giralt, Mayrhofer and Sheard1995; Sheard Reference Sheard, Nash, Ryan, Diederich, Gries and Bungartz2004). Finally, R. placynthielloides, known only from the type locality in Taiwan, has small lecideine apothecia, up to 0·5 mm diam., smaller Pachysporaria-type ascospores up to 20 × 10 μm, 4-spored asci and lacks secondary metabolites (Aptroot & Sparrius Reference Aptroot and Sparrius2003; see also Giralt & van den Boom Reference Giralt and van den Boom2008).
There are other vegetatively reproducing Rinodina species but they are all blastidiate or sorediate instead of isidiate and therefore cannot be mistaken for R. fuscoisidiata. The main studies including Rinodina taxa developing vegetative propagules are the following: Malme (Reference Malme1902), Sheard (Reference Sheard1995) and Giralt et al. (Reference Giralt, Mayrhofer and Sheard1995), for the corticolous species; and Matzer & Mayrhofer (Reference Matzer and Mayrhofer1994, Reference Matzer and Mayrhofer1996) and Giralt & van den Boom (Reference Giralt and van den Boom2008), for the saxicolous species. Additional blastidiate or sorediate Rinodina species not included in the compilations mentioned above are: R. pityrea Ropin & H. Mayrhofer (Ropin & Mayrhofer Reference Ropin and Mayrhofer1995); R. australiensis Müll. Arg. (Mayrhofer et al. Reference Mayrhofer, Kantvilas and Ropin1999); R. perreagens Sheard and R. juniperina Sheard (Sheard & Mayrhofer Reference Sheard and Mayrhofer2002; Sheard Reference Sheard, Nash, Ryan, Diederich, Gries and Bungartz2004); R. turfaceoides van den Boom et al. (Giralt et al. Reference Giralt, Mayrhofer, van den Boom and Elix2001); and R. evae Fos & Giralt (Fos & Giralt Reference Fos and Giralt2009).
Additional specimens examined. Venezuela: Mérida: distr. Rangel, zwischen Laguna Mucubaji und Pico Mucuñuque, etwa 15 km SE von Apartaderos, in Paramo-Vegetation, 3500 m, 8°45′N; 70°45′W, 1989, K. & A. Kalb & López-Figueiras (hb. Kalb 27015—topotype); Sierra de Santo Domingo, umbegung des Karsees Laguna Negra, 3500 m, 1969, B. & F. Oberwinkler & H. Hertel 10474 (M); 3 km NE of Pico El Águila, 3900–4000 m, Paramo, on an open rocky slope, 1979, R. Santesson 29442 (UPS); distr. Miranda, Paramo zwischen Almorzadero und Piñango, 4200 m, 8°55′N; 70°50′W, 1989, K & A. Kalb (hb. Kalb-27016).
The authors are indebted to the curators of the herbaria BCN, M and UPS, and to Helmut Mayrhofer for providing valuable information on additional specimens. The first author also thanks the ‘Comissionat per a la Recerca’ (Catalan Government) and the project CGL2007-66734-C03-02/BOS (Spanish Government) for financial support.