Introduction
Lepraria Ach. is a cosmopolitan genus of leprose to byssoid, permanently sterile lichenized fungi, with more than 60 species known worldwide (e.g. Laundon Reference Laundon1989, Reference Laundon1992; Tønsberg Reference Tønsberg1992, Reference Tønsberg2007; Sipman Reference Sipman2004; Slavíková-Bayerová & Orange Reference Slavíková-Bayerová and Orange2006; Flakus & Kukwa Reference Flakus and Kukwa2007; Knudsen & Elix Reference Knudsen and Elix2007a; Slavíková-Bayerová & Fehrer Reference Slavíková-Bayerová and Fehrer2007; Elix & Kalb Reference Elix and Kalb2008; Lendemer et al. Reference Lendemer, Knudsen and Elix2008; Kukwa & Flakus Reference Kukwa and Flakus2009; Saag et al. Reference Saag, Saag and Randlane2009). Of these, 22 species have been reported from South America (see Calvelo & Liberatore Reference Calvelo and Liberatore2002; Sipman Reference Sipman2004; Flakus et al. Reference Flakus, Kukwa and Czarnota2006; Flakus & Kukwa Reference Flakus and Kukwa2007; Nöske et al. Reference Nöske, Mandl and Sipman2007; Kukwa & Flakus Reference Kukwa and Flakus2009 and literature cited therein). However, the number of Lepraria present in the continent is expected to be much greater. In this paper we present new records of 18 Lepraria species, two of them new to the Southern Hemisphere and one new to South America, with the description of two new species from Argentina and Bolivia. As a consequence, the number of Lepraria known from South America is enlarged to 27 species. In addition 4-O-methylleprolomin is discovered in lichens for the second time from a new species of Lepraria, and is new to Lepraria. Most of the taxa presented here were discussed previously by Flakus & Kukwa (Reference Flakus and Kukwa2007) and Saag et al. (Reference Saag, Saag and Randlane2009), so we present only a brief mention of the characteristics of each species.
Material and Methods
Most of the examined specimens were collected by the authors in Bolivia and Peru in 2008, and these are deposited in KRAM, LPB and UGDA, with duplicates donated to B, BG, CANB, NY, UCR and hb. Flakus. Additional specimens studied were requested on loan from B, H, PRA and S. The abbreviations AF, MK and PR in specimens examined stand for A. Flakus, M. Kukwa and P. Rodriguez, respectively.
Lichen morphology and anatomy were examined using standard stereo- and compound microscopes. Squash preparations were studied in water or KOH solution (K), with or without pre-treatment with acetone. The granules and crystals covering hyphae of L. nothofagi were also studied in polarized light. The measurements of soredia were made on material mounted in water.
Chemical analyses were carried out using thin-layer chromatography (TLC) following the methods of Orange et al. (Reference Orange, James and White2001). Extracts were run on aluminium or glass TLC plates in solvent systems A, B and C. Confirmation of identified substances was achieved in some cases by running the extracts adjacent to an extract containing known substances. Some specimens of newly described taxa were analysed by HPLC. When the substance was invariably present it is marked with (+), but if the metabolite was absent from some specimens (±) is used.
Results
Lepraria achariana Flakus & Kukwa
Chemistry. Lecanoric (+) and angardianic/roccellic (±) acids.
Notes. Known in South America only from Bolivia; here we report a fifth locality. The specimen lacked anthraquinones, which were reported for this species by Flakus & Kukwa (Reference Flakus and Kukwa2007).
Specimens examined. Bolivia: Dept. Cochabamba: Prov. Carrasco, Carrasco National Park, between Sehuencas and Monte Punku villages, alt. c. 2650 m, MK 6572, 6573 & 6582a (LPB, UGDA).
Lepraria adhaerens K. Knudsen, Elix & Lendemer
Chemistry. Pannarin (+) and zeorin (+), as well as two unidentified substances (visible in long UV wavelength as ice-blue spots before charring, and after charring on fresh plates) (+) in Rf class: A7, B7, C6–7 and A5, B4, C5.
Notes. For the characteristics of L. adhaerens see Knudsen et al. (Reference Knudsen, Elix and Lendemer2007). Our specimens are morphologically and chemically (pannarin and zeorin present) very similar to the isotype deposited in UGDA, so we have attributed them to L. adhaerens. However, they always produce two additional unidentified substances (see above), so far not recorded for this species or any other member of the genus. The taxonomic significance of these unknown metabolites is not certain and needs further study on more extensive collections. The South American collections may eventually represent an undescribed taxon, but this should be clarified by using molecular methods.
Lepraria santamonicae K. Knudsen & Elix is morphologically similar to L. adhaerens, but is greener in colour and contains argopsin and norargopsin. So far it has not been reported outside North America (Knudsen & Elix Reference Knudsen and Elix2007a; Saag et al. Reference Saag, Saag and Randlane2009).
Previously L. adhaerens has been reported only from North America (Pennsylvania, Missouri, California), where it grows on soil and acid or limestone rocks (Knudsen et al. Reference Knudsen, Elix and Lendemer2007; Knudsen & Elix Reference Knudsen, Elix, Nash, Gries and Bungartz2007b). Here we record this species for the first time from the Southern Hemisphere. Peruvian specimens were found on soil, terricolous bryophytes, rocks, and sometimes on small twigs in an open semi-desert montane area.
Specimens examined. Peru: Dept. Arequipa: Prov. Caylloma, Cañon del Colca canyon, Llahuar village, alt. 2085 m, 15°34′52″S, 72°00′37″W, AF 10143 (KRAM), MK 6115 & 6118 (UGDA); between Soro and Llahuar villages, alt. 2100 m, 15°34′41″S, 72°01′01″W, AF 10130, 10137 (KRAM, hb. Flakus) & MK 6102 & 6109 (UGDA); Soro village, alt. 2050 m, 15°34′50″S, 72°01′51″W, MK 6136 (UGDA).
Lepraria alpina (B. de Lesd.) Tretiach & Baruffo
Chemistry. TLC: atranorin (+), porphyrilic acid (+) and an unknown substance below porphyrylic acid (±). No fatty acids were found in the material studied, but they have been reported by Flakus & Kukwa (Reference Flakus and Kukwa2007).
Notes. For the descriptions of L. alpina see Tønsberg (Reference Tønsberg1992), Leuckert et al. (Reference Leuckert, Kümmerling and Wirth1995) and Baruffo et al. (Reference Baruffo, Zedda, Elix and Tretiach2006). Most specimens of L. alpina from South America that we examined are characterized by a whiter and powdery thallus surface (for more details, see Flakus & Kukwa Reference Flakus and Kukwa2007). The status of this material from South America is not clear and can be resolved only by molecular studies.
In South America L. alpina was previously known only from Bolivia. Here the species is reported as new to Chile, Colombia, Peru and Venezuela. It is mainly a saxicolous (on sandstone, siliceous or basic rocks) and muscicolous species, but rarely occurs on bark.
Specimens examined. Chile: Terr. Magellanes: Lago del Toro (L. Maravilla), Estancia Rio Payne, above the river, Santesson 6579 (S–L42671); Natales, Cerro Dorotea, Santesson 2183 & 8260 (S–L42669 & L42670).—Colombia: Risaralda: Mnpio. de Sta. Rosa de Cabal, Camino real entre Termales de Sta. Rosa y Hda. La Sierra, 4°50′N, 75°31′W, alt. 3190 m, Wolf 981 (B).—Peru: Dept. Arequipa: Prov. Caylloma, by road to Huambo village near Lagunillas pass, alt. 4240 m, 15°55′10″S, 72°09′12″W, AF 10128 (KRAM) & MK 6092 (UGDA).—Venezuela: Sierra Nevada: Edo. Merida, Mun. Libertador, in 4105 m Höhe, Wiegleb 580 (B). Region de Araucaria: Mininco, zwischen Gestein, trocken Stellen, häufig, Kunkel 2074 (B).
Lepraria borealis Lohtander & Tønsberg
Chemistry. Atranorin (+), rangiformic (+) and norrangiformic acids (+).
Notes. Lepraria borealis is morphologically very similar to other members of the L. neglecta group (particularly to L. caesioalba), but Ekman and Tønsberg (Reference Ekman and Tønsberg2002) confirmed that it is a distinct species. For descriptions of the species see Tønsberg (Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004), Kukwa (Reference Kukwa2006a), Slaviková-Bayerová & Fehrer (2007) and Saag et al. (Reference Saag, Saag and Randlane2009).
Previously L. borealis was known from Europe, North America including Greenland, Antarctica and Africa (Canary Islands) (Tønsberg Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004; Baruffo et al. Reference Baruffo, Zedda, Elix and Tretiach2006; Kukwa Reference Kukwa2006a; Saag et al. Reference Saag, Saag and Randlane2009). Here the species is reported for the first time from South America. The Chilean specimen was collected on wood.
Specimen examined. Chile: Ñuble: Chillán (Bureo), Barros 106 (H).
Lepraria caesioalba (B. de Lesd.) J. R. Laundon
Chemistry. The South American specimens are represented by chemotype I (see Leuckert et al. Reference Leuckert, Kümmerling and Wirth1995) containing fumarprotocetraric (+) acid and atranorin (+), and chemotype III with atranorin (+), psoromic (+), and conpsoromic acids (+), with accessory roccellic, rangiformic and/or unknown fatty acids.
Notes. Lepraria caesioalba s. lat. was thought to be a paraphyletic species (see Ekman & Tønsberg Reference Ekman and Tønsberg2002; Slaviková-Bayerová & Fehrer 2007). It was represented by several chemotypes (see Leuckert et al. Reference Leuckert, Kümmerling and Wirth1995; Tønsberg Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004 and literature cited therein), of which the chemotype with stictic acid has been recently segregated as L. svalbardensis Tønsberg (Øvstedal et al. Reference Leuckert, Kümmerling and Wirth2009); other chemotypes may also represent separate species. Amongst the South American material we encountered chemotypes I and III sensu Leuckert et al. (Reference Leuckert, Kümmerling and Wirth1995), which are phylogenetically closely related and may represent one chemically variable species (see Ekman & Tønsberg Reference Ekman and Tønsberg2002). See also Flakus & Kukwa (Reference Flakus and Kukwa2007).
In South America L. caesioalba is known from Argentina, Bolivia, Colombia and Peru (Sipman Reference Sipman2004; Flakus & Kukwa Reference Flakus and Kukwa2007; Sipman et al. Reference Sipman, Hekking and Aguirre2008), and appears to be rather common in open, high montane areas. Here it is reported as new to Venezuela where it was found on soil, rock and terricolous bryophytes in Andean vegetation.
Specimens examined. Argentina: South-West Patagonia: Lago Frio, western side, James 1240 (BM). Tierra del Fuego: Sierra Alvear, the southern slope, above Las Cotorras (about 20 km ENE of Ushuaia), c. 600 m, Santesson 774 (S–L42673; chemotype I).—Peru: Prov. Canchis: Sibinasallma village, Murmurani sector, W of la Laguna Sibinacocha, alt. 4952 m, 13°50′05″S, 71°04′06″W, PR 567 (LPB; chemotype III).—Venezuela: Sierra Nevada: Edo. Merida, Mun. Liberador, unterhalb des Pico Espejo, Weg Loma Redonda – Pico Espejo, alt. 4440 m, Wiegleb 622a (B; chemotype I).
Lepraria diffusa (J. R. Laundon) Kukwa
Chemistry. Atranorin (±), 4-oxypannaric acid 2-methyl ester (+), pannaric acid (+), angardianic/roccellic acid (±) and traces of unknowns (+).
Notes. This species is characterized by the production of 4-oxypannaric acid-2-methyl ester and related substances (for more detail see Laundon Reference Laundon1989; Elix & Tønsberg Reference Elix and Tønsberg2004; Tønsberg Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004; Kukwa Reference Kukwa2006a; Saag et al. Reference Saag, Saag and Randlane2009). In South America L. diffusa can be confused with the morphologically and chemically similar L. vouauxii. The two species can readily be distinguished chemically, as L. vouauxii contains high concentrations of pannaric acid 6-methyl ester (Tønsberg Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004; Kukwa Reference Kukwa2006a).
Previously this species was known from Europe, North America and Asia (e.g., Tønsberg Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004; Saag et al. Reference Saag, Saag and Randlane2009). Here it is recorded for the first time from the Southern Hemisphere, from Bolivia, Ecuador and Peru, where it was collected from soil in the Andes, mainly in cloud forests and Páramo vegetation.
Specimens examined. Bolivia: Dept. Cochabamba: Prov. Carrasco, Carrasco National Park, Monte Punku village, alt. 2800 m, 17°35′01″S, 65°18′03″W, MK 6152 & 6156 (LPB, UGDA); between Sehuencas and Monte Punku villages, alt. c. 2650 m, MK 6572 (LPB, UGDA); near Sehuencas village by Rio Lopez Mendoza river, alt. 2226 m, 17°30′26″S, 65°16′55″W, MK 6590 (LPB, UGDA).—Chile: X Region: Parque Nacional Puyehue, Anticura, Los Derrumbes, 40°40′S, 72°10′W, alt. c. 600 m, Coppins, Galloway, Guzmán & James 4643 (BM; mixed in specimen of L. simpaniana).—Ecuador: Prov. Pichincha: Volcán Pichincha, E slope, along road to antennas at summit, 00°10′S, 78°35′W, 3200–3700 m, Harris 17054 (B); Prov. Carchi: Volcán Chíles, at road-side between Maldonado and Tufiño above the lake Laguna Verde, c. 1·5–1·8 km SSE of the peak of Volcán Chíles, alt. 4050 m, 00°48′ N, 77°55′30″ W, Etayo & Palice 8357 (PRA).—Peru: Dept. Junin: Prov. Tarma, c. 10 km (road distance) NNE of Palca, c. 2600 m 11°18′S, 75°32′W, Santesson P12:60 & Moberg (S–L42642).
Lepraria ecorticata (J. R. Laundon) Kukwa
Chemistry. Usnic acid (+), zeorin (+) and trace of unidentified substance (±).
Notes. The specimens cited below fall well within the variation exhibited by L. ecorticata (for the characteristics of the species see Laundon Reference Laundon2003; Kukwa Reference Kukwa2006b; Flakus & Kukwa Reference Flakus and Kukwa2007), but we have also seen specimens (not cited here) with the same chemistry but different morphology. One saxicolous Bolivian specimen consisted of soredia dispersed over a very thin layer of prothallus; the soredia were fluffy and possessed short projecting hyphae. Four specimens, three from Peru and one from Bolivia, had irregularly shaped granules and were morphologically very similar to L. texta K. Knudsen, Elix & Lendemer (see Knudsen & Elix Reference Knudsen, Elix, Nash, Gries and Bungartz2007b), but lacked atranorin and fatty acids. The identification of those specimens no doubt requires a molecular approach.
Recently the phylogenetic position of two Lepraria species containing usnic acid and zeorin, L. coriensis (Hue) Sipman and L. usnica Sipman, was studied and it was shown that they do not belong to Lepraria s. str. (Nelsen et al. Reference Nelsen, Lumbsch, Lücking and Elix2008). This is also possibly the case with L. ecorticata and with L. leuckertiana, treated below.
In South America L. ecorticata has been reported from Bolivia, Brazil, Chile and Peru (Kukwa Reference Kukwa2006b; Flakus & Kukwa Reference Flakus and Kukwa2007). In this paper we present new records from Bolivia and Peru. Usually this species occurs on rocks, saxicolous bryophytes and moribund lichens in high, open Andean vegetation and montane cloud forests, and on siliceous rocks in lowland Amazon forest.
Specimens examined. Bolivia: Dept. La Paz: Prov. Nor Yungas, near Pacallo village, alt. 1360 m, 16°12′10″S, 67°50′39″W, AF 11820 & MK (KRAM, LPB).—Peru: Dept. Cuzco: Prov. Urubamba, Malaga Chico (Tastiyiocc), NNW of Ollantaytambo, alt. 4000–4100 m, 13°04′S, 72°22′W, Santesson P93:122 & Moberg (S–L42651).
Lepraria incana (L.) Ach
Chemistry. Divaricatic acid (+) and zeorin (+) were detected in the specimen studied.
Notes. For the characteristics of this common species and the differentiation with other taxa see Tønsberg (Reference Tønsberg1992), van den Boom et al. (1994), Aptroot et al. (Reference Aptroot, Diederich, Sérusiaux and Sipman1997) and Kukwa (Reference Kukwa2006a).
Lepraria incana is a rather widespread species, but in South America it was previously known only from Bolivia and Colombia (Flakus & Kukwa Reference Flakus and Kukwa2007; Sipman et al. Reference Sipman, Hekking and Aguirre2008); here it is reported for the first time for Chile and from a second locality in Bolivia. The South American material occurred on tree ferns, epiphytic bryophytes or Parnettya twigs.
Specimens examined. Bolivia: Dept. Cochabamba: Prov. Carrasco, Carrasco National Park, near Sehuencas village, alt. 2220 m, 17°30′12″S, 65°16′30″W, MK 6559 (LPB, UGDA).—Chile: Juan Fernandez Islands: Masatierra, Pangal, 500 m, Kunkel 305/4 (B).
Lepraria lecanorica Tønsberg
Chemistry. Atranorin (+) and lecanoric acid (+).
Notes. For a description of the species see Tønsberg (Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004).
Lepraria lecanorica is known from North and South America (Tønsberg Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004; Flakus & Kukwa Reference Flakus and Kukwa2007). Here the reported locality is the third for South America and the second for Chile.
Specimen examined. Chile: Prov. Coquimbo: Depto. Ovalle, Fray Jorge, alt. 300 m, Sparre 3087 (S).
Lepraria leuckertiana (Zedda) L. Saag
Chemistry. Usnic acid (+) and zeorin (+).
Notes. For the characteristics of the species see Zedda (Reference Zedda2000), Flakus et al. (Reference Flakus, Kukwa and Czarnota2006) and Saag et al. (Reference Saag, Saag and Randlane2009).
Lepraria leuckertiana is known mainly from southern Europe and north Africa, but it has also recently been found in South America (Zedda Reference Zedda2000; Flakus et al. Reference Flakus, Kukwa and Czarnota2006; Saag et al. Reference Saag, Saag and Randlane2009 and literature cited therein). Here we report a second South American locality for this species from Peru, where it grew on soil.
Specimen examined. Peru: Dept. Pasco: Prov. Cerro de Pasco, c. 7 km NE of Cerro de Pasco, alt. c. 3900 m, 10°40′S, 76°14′W, Santesson P29:5 & Moberg (S).
Lepraria lobificans Nyl
Chemistry. Atranorin (+), zeorin (+) and the stictic acid complex (+). In some specimens a pink spot due to an unknown substance (±) was sometimes found below zeorin.
Notes. For the characteristics of the species see Tønsberg (Reference Tønsberg1992, 2004) and Flakus & Kukwa (Reference Flakus and Kukwa2007).
Lepraria lobificans is a common species in South America, especially in temperate and colder regions (e.g. Sipman Reference Sipman2004; Flakus & Kukwa Reference Flakus and Kukwa2007). According to our knowledge, this is the first report of this species from Argentina. The specimens listed here were collected on rock, tree bark, soil and saxicolous bryophytes in forests and semi-desert montane areas.
Specimens examined. Argentina: Prov. Tucuman: Dep. Monteros, Rio de los Sosas, alt. 1000 m, Digilio-Grassi 912 (S).—Bolivia: Dept. Cochabamba: Prov. Carrasco, Carrasco N. P., near Sehuencas village, alt. 2220 m, 17°30′12″S, 65°16′30″W, AF 10479, 10507, 10551, MK 6418 & 6463 & PR (KRAM, LPB, UGDA); 18 km N of Monte Punku village, between Ch'iqta rumi and Phaqcha settlements, alt. 2700 m, 17°27′22″S, 65°16′24″W, AF 10242, MK 6256, 6263 & PR (KRAM, LPB, UGDA); Prov. Nor Yungas, near Pacallo village, alt. 1360 m, 16°12′10″S, 67°50′39″W, AF 11820/1, 11815, 11818, 11828, 11835 & MK 7130, 7133, 7134, 7143, 7145 (KRAM, LPB, UGDA, herb. Flakus).—Chile: Prov. Valdivia: Lago Riñihue, Riñihue, Cerro Tralcan, Santesson 3809 (S). Juan Fernandez Islands: Caves of the Expelled, near Bahia de Cumberland, Kunkel 336a/22 (B).—Peru: Dept. Arequipa: Prov. Caylloma, Cañon del Colca canyon, between Soro and Llahuar villages, alt. 2100 m, 15°34′41″S, 72°01′01″W, MK 6099 (UGDA).
Lepraria nothofagi Elix & Kukwa sp. nov
Similis speciei Lepraria lecanorica, sed acidum porphyrilicum et strepsilinum continente differt.
Typus: Argentina, Gobernación del Neuquén, Lago Quillén, on point by the easternmost bend of the lake, on level ground on N shore, Nothofagus obliqua forest, on bark of Nothofagus, 28 December 1937, A. Kalela 173b (H—holotypus; UGDA—isotypus).
Thallus white to ivory white, in one specimen with a yellowish tinge in part, delimited to obscurely lobed, when young consisting of aggregations of ± loose soredia; medulla absent or present in older thalli, thick, but perhaps consisting only of old and bleached soredia, as some algal cells were found among hyphae; hypothallus of sparse hyphae attaching the thallus to the substratum, hyphae up to 3·5 µm wide, branched, rarely anastomosing, bare or covered with granules (not apparent in polarized light, insoluble in K, N and acetone) and crystals (up to c. 10 × 5 µm, white in polarized light, insoluble in K and acetone, but soluble in N); soredia simple or forming consoredia, free-laying or embedded in the medulla; soredia up to 90(–150) µm diam., consoredia up to c. 180(–250) µm diam.; projecting hyphae present, up to 15(–40) µm long, or absent; soredial wall not complete; photobiont chlorococcoid, up to 10 µm diam.
Chemistry. Atranorin (+; major), strepsilin (+; major to minor), porphyrilic acid (+; major to minor) and lecanoric acid (+; trace) were detectable by TLC. Two specimens were studied by HPLC, and atranorin (+; major), strepsilin (+; major to minor), porphyrilic acid (+; major to minor), lecanoric acid (+; trace), norascomatic acid (±; minor), di-O-methylstrepsilin (±; trace), pannaric acid (±; minor) were found. Some soredia reacted C+ green due to the presence of strepsilin.
Notes. This new species is characterized by the white, thick, obscurely lobate and stratified thallus and the production of atranorin, strepsilin, porphyrilic acids and a constant trace of lecanoric acid. Morphologically L. nothofagi closely resembles L. lecanorica, but the latter contains lecanoric acid in high concentrations and lacks strepsilin and porphyrilic acid (Tønsberg Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004). Chemically, this new species is similar to L. xerophila Tønsberg, especially to chemotype II due to the presence of strepsilin. That chemotype, however, produces norascomatic acid as a major secondary metabolite, lacks porphyrilic and lecanoric acids, and strepsilin occurs always as a minor compound. In chemotype I of L. xerophila, pannaric acid 6-methyl ester is always present as the major secondary substance, sometimes together with porphyrilic and pannaric acids (Elix & Tønsberg Reference Elix and Tønsberg2004; Tønsberg Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004). Additionally, the morphology of L. xerophila is very different; the thallus can be squamulose to crustose with an effigurate to placodioid margin, the thallus surface bears numerous regular to irregular and deformed, fragile, more or less dorsiventral (sometimes capitate) easily detached lobules which act as diaspores, and a distinct algal layer is present in the thallus (Tønsberg Reference Tønsberg, Nash, Ryan, Diederich, Gries and Bungartz2004).
Strepsilin also occurs in L. dibenzofuranica Elix, L. goughensis Elix & Øvstedal and L. multiacida Aptroot (Elix & Tønsberg Reference Elix and Tønsberg2004; Elix et al. Reference Elix, Øvstedal and Gremmen2005; Elix & Kalb Reference Elix and Kalb2008). Lepraria dibenzofuranica has a powdery, whitish green to greenish or bluish grey thallus containing several dibenzofurans (strepsilin in minor or trace amounts), anthraquinones and fatty acids (Elix & Kalb Reference Elix and Kalb2008). Lepraria goughensis produces lecanoric and gyrophoric acids and anthraquinones in addition to strepsilin and has a powdery thallus but lacks a hypothallus and medulla (Elix et al. Reference Elix, Øvstedal and Gremmen2005). Lepraria multiacida differs in having a stratified and sometimes lobate thallus containing atranorin, the stictic acid complex, consalazinic and salazinic acids, terpenoids, and some other substances in addition to strepsilin (Aptroot Reference Aptroot2002; Elix & Tønsberg Reference Elix and Tønsberg2004).
Porphyrilic acid is more commonly produced than strepsilin by various Lepraria species (Elix & Tønsberg Reference Elix and Tønsberg2004; Kukwa & Flakus Reference Kukwa and Flakus2009; Saag et al. Reference Saag, Saag and Randlane2009), and in all cases, the taxa differ significantly in chemistry. In South America two taxa are known to produce porphyrilic acid: L. alpina and L. neojackii Flakus & Kukwa (Flakus & Kukwa Reference Flakus and Kukwa2007). The first species belongs to the L. neglecta group and has a coarsely granular thallus, often produces fatty acids and lacks strepsilin (see under that species above). Lepraria neojackii has a green-grey, leprose thallus containing porphyrilic acid, fatty acids, anthraquinones but lacks atranorin (Flakus & Kukwa Reference Flakus and Kukwa2007).
At present this species is known from several locations near the type locality in Argentina. It has always been found on the bark of Nothofagus in forest dominated by Austrocedrus or Nothofagus species (N. dombeyi, N. obliqua or N. pumiloo).
Specimens examined (all paratypes). Argentina: Gobernacion del Neuquen: Lago Traful, steep N slope near lake shore, c. 2 km W of the school, alt. 854 m, Kalela 52a (H); Lago Lacar, on dry E slope N of San Martin, Kalela 197b (H); Lago Quillén, on gentle N slope of S shore of the lake, Kalela 178 (H); Lago Quillén, N shore of the lake, Kalela 171b (H); Lago Quillén, S of slope of hill at the eastern end of the lake, A. Kalela 181b:c (H); Lago Quillén, by the eastern bend of the lake, A. Kalela 174e, 177b & 180c (H); Lago Quillén, on level ground on rock at the E end of the lake, A. Kalela 187a (H). Lago Traful, steep slope N of the school, A. Kalela 50d (H, UGDA); Lago Correntoso, on top of little height on the isthmus between Lago Correntoso and Lago Espejo, A. Kalela 29h (H, UGDA).
Lepraria pallida Sipman
Chemistry. Atranorin (+), zeorin (+) and two unidentified fatty acids (+).
Notes. For the characteristics of the species see Sipman (Reference Sipman2004). Some of the Bolivian specimens appeared juvenile, with only obscurely lobed thalli and a poorly developed hypothallus, being composed mainly of white to grey hyphae. Such material was usually mixed with typical, well-developed specimens.
This species has been reported from South America (Bolivia, Brazil), Madagascar, the Seychelles and Asia (Sipman Reference Sipman2004; Flakus & Kukwa Reference Flakus and Kukwa2007; Kukwa & Flakus Reference Kukwa and Flakus2009). Here it is recorded for the first time from Peru. Specimens cited below were collected on rock, soil, tree bark and saxicolous bryophytes in montane forests.
Specimens examined. Bolivia: Dept. Cochabamba: Prov. Carrasco, Carrasco N. P., near Sehuencas village, alt. 2220 m, 17°30′12″S, 65°16′30″W, AF 10387, 10469, 10471, 10473, MK 6462, 6496, 6543 & PR (KRAM, LPB, UGDA); Dept. La Paz: Prov. Nor Yungas, near Pacallo village, alt. 1360 m, 16°12′10″S, 67°50′39″W, AF 11805, 11814 & MK (KRAM, LPB, hb. Flakus).—Peru: Dept. San Martin: Prov. San Martin, Cerro Escalera (NE of Tarapoto), NW of the tunnel, alt. c. 1000 m, 06°26′S, 76°15′W, Santesson P74:91 & Thor (S).
Lepraria sipmaniana (Kümmerling & Leuckert) Kukwa
Chemistry. Pannaric acid 6-methylester (+), 4-oxypannaric acid 6-methylester (+), anthraquinones (±), ‘vouauxii unknown 1’ (±) and traces of unknowns (±).
Notes. For the characteristics of the species see Leuckert & Kümmerling (Reference Leuckert and Kümmerling1991), Sipman (Reference Sipman2004) and Flakus & Kukwa (Reference Flakus and Kukwa2007).
Recently we revised the specimens of L. membranacea (Dicks.) Vain. reported from Chile by Laundon (Reference Laundon1989). They are indeed morphologically similar to L. membranacea, but contain pannaric acid 6-methylester as the major dibenzofuran, have distinctly raised lobe margins and possess granules dispersed over the surface; this latter characteristic is typical of L. sipmaniana, described several years after Laundon's monograph on the L. membranacea group (see Leuckert & Kümmerling Reference Leuckert and Kümmerling1991). As a consequence, the Chilean records of L. membranacea are considered to be erroneous.
Lepraria membranacea has also been reported from Argentina (Calvelo & Liberatore Reference Calvelo and Liberatore2002). However, we have not seen this material, and given the distribution of L. sipmaniana and L. membranacea, the Argentinian records of the latter should be considered as doubtful.
In South America L. sipmaniana has been recorded from Brazil, Colombia and Peru (Leuckert & Kümmerling Reference Leuckert and Kümmerling1991; Flakus & Kukwa Reference Flakus and Kukwa2007). Here it is reported as new to Bolivia and Chile. In the latter localities it grew on soil, rocks and terricolous bryophytes in open, high Andean, semi-desert areas or in montane forests.
Specimens examined. Bolivia: Dept. Cochabamba: Prov. Carrasco, Carrasco N. P., near Sehuencas village, alt. 2220 m, 17°30′12″S, 65°16′30″W, MK 6479 (LPB, UGDA).—Brazil: Rio de Janeiro: Restinga südlich der Stadt, Mattick 1613 (B).—Chile: IX Region: La Araucania, 5 km NE of Melipeuco, alt. 400–600 m, 38°49′S, 71°40′W, Coppins, Galloway, Guzmán & James 5635 (BM; as Leproloma membranaceum). X Region: Parque Nacional Puyehue, Anticura, Los Derrumbes, 40°40′S, 72°10′W, alt. c. 600 m, Coppins, Galloway, Guzmán & James 4643 (BM; as L. membranaceum).—Peru: Dept. Arequipa: Prov. Caylloma, below Lagunillas pass, alt. 4200 m, 15°54′54″S, 72°10′12″W, AF 10132 & MK (KRAM); Prov. Caylloma, Cañon del Colca canyon, between Soro and Llahuar villages, alt. 2100 m, 15°34′41″S, 72°01′01″W, MK 6097 & 6098 (UGDA). Dept. Huanuco: Prov. Huanuco, valley of Rio Higueras, c. 17 km (road distance) W of Huanuco, alt. 2300 m, 09°55′S, 76°22′W, Santesson P48:9 & Moberg (S). Dept. Junin: Prov. Tarma, c. 10 km (road distance) NNE of Palca, c. 2600 m, 11°18′S, 75°32′W, Santesson P12:60 & Moberg (S, mixed in specimen of L. diffusa).
Lepraria squamatica Elix
Chemistry. Squamatic (+) and baeomycetic (+) acids.
Notes. For a description of the species see Elix (Reference Elix2006).
Previously L. squamatica was only known from scattered localities in Australia (Elix Reference Elix2006, 2009), South America (Bolivia) (Flakus & Kukwa Reference Flakus and Kukwa2007) and North America (Lendemer Reference Lendemer2008). Here it is reported as new to Peru.
Specimen examined. Peru: Dept. San Martin: Prov. Lamas, Cerro Blanco, c. 58 km W-WNW of Tarapoto (road distance), c. 06°25′S, 76°40′W, c. 1000 m, Santesson P73:41 & Thor (S).
Lepraria stephaniana Elix, Flakus & Kukwa sp. nov
Thallus leprosus, virido- vel flavido-albidus, plerumque non stratosus, elobatus, soredia crassa, cum hyphis filamentosis projectis; acidum salazinicum, zeorinum, 4-O-methylleprolominum et terpenum ignotus (Rf A6, B6, C6) continens.
Typus: Bolivia, Dept. La Paz, Prov. Iturralde, between Ixiamas and Santa Rosa de Maravillas villages, alt. 305 m, 13°49′16″S, 68°07′18″W, preandean Amazon forest, on bark of tree, 28 July 2008, M. Kukwa 6828 (UGDA-L—holotypus; B, BG, KRAM, LPB, NY, UCR—isotypi).
Paratypes: Bolivia: Dept. La Paz: locality as for the holotype, 28 July 2008, A. Flakus 11192 & M. Kukwa 6852 (CANB, KRAM, UGDA, LPB, herb. Flakus).
Thallus crustose, thick, usually not delimited nor lobed, green-grey to creamy white, not stratified, but sometimes with a poorly differentiated, pseudo-medullary layer of decaying soredia; hypothallus indistinct; soredia irregularly spreading over the substratum, coarse with soft appearance, irregularly rounded, up to 100(–200) µm diam., composed of very lax hyphae mixed with algal cells, usually with projecting hyphae up to c. 30(–50) µm long; soredial wall incomplete; photobiont chlorococcoid, cells up to 12 µm diam.
Chemistry. 4-O-methylleprolomin (major), salazinic acid (minor), zeorin (minor) and an unknown terpenoid (minor) with Rf class values A6, B6, C6 were detected in all specimens. Thallus reacts K+ yellow turning brownish to red, P+ yellow, C−, KC−.
Etymology. The new species is named in honour of the prominent German botanist, Dr. Stephan G. Beck (La Paz), the founder of Herbario Nacional de Bolivia in La Paz, for his substantial contributions to the knowledge of the Bolivian flora.
Notes. Lepraria stephaniana is a very distinctive species characterized by the thick, unstratified and non-lobed thallus composed of coarse soredia with soft appearance, and unique chemistry. This is the first species of the genus known to produce 4-O-methylleprolomin as a major secondary substance. This unique diphenyl ether has been previously detected in a Pannaria species (A. Elvebakk & J. A. Elix, unpublished).
In addition to L. stephaniana, salazinic acid is only known from two other Lepraria species, namely L. multiacida and L. salazinica Tønsberg (Aptroot Reference Aptroot2002; Elix & Tønsberg Reference Elix and Tønsberg2004; Tønsberg Reference Tønsberg2007). Lepraria multiacida can be clearly distinguished by its stratified, often lobate thallus containing numerous substances (stictic, norstictic, connorstictic and consalazinic acids, atranorin and strepsilin) not present in L. stephaniana (Aptroot Reference Aptroot2002; Elix & Tønsberg Reference Elix and Tønsberg2004). Lepraria salazinica, known only from North America, has smaller soredia (up to 30 µm in diam.) than L. stephaniana, and contains additional atranorin and angardianic/roccellic acid (Tønsberg Reference Tønsberg2007) rather than 4-O-methylleprolomin and terpenoids.
The new species has been found in preAndean Amazon forest in Bolivia, where large populations were observed on the bark of two trees. So far it is known only from the type locality.
Lepraria vouauxii (Hue) R. C. Harris
Chemistry. Pannaric acid 6-methylester (+), 4-oxypannaric acid 6-methylester (+), ‘vouauxii unknown 1’ (±, see Tønsberg Reference Tønsberg1992), atranorin (±) and traces of unknowns (±).
Notes. For descriptions of the species see Elix & Tønsberg (Reference Elix and Tønsberg2004), Flakus & Kukwa (Reference Flakus and Kukwa2007) and Saag et al. (Reference Saag, Saag and Randlane2009).
Lepraria vouauxii is a widely distributed, but still under-recorded species known from various substrata (see e.g. Laundon Reference Laundon1989; Leuckert & Kümmerling Reference Leuckert and Kümmerling1991; Tønsberg Reference Tønsberg1992, 2004; Kukwa & Flakus Reference Flakus and Kukwa2007, 2009; Elix Reference Elix2009). Here new records from Bolivia, Chile, Ecuador and Peru are added.
Specimens examined. Bolivia: Dept. La Paz: Prov. Franz Tamayo, W of La Paz city near Ulla Ulla village, Mt. Puntani, PR 565 (LPB).—Chile: Terr. Magallanes: Lago del Toro (L. Maravilla), Estancia Rio Payne, above the river, Santesson 6594 (S).—Ecuador: Prov. León: Railway station Cotopaxi, alt. 3550 m, Asplund L63 (S); Prov. Pichincha: c. 25 km N of Quito, S-side of Pululagua caldera, alt. 2750–2800 m, Palice 3761 & Soldán (PRA); Prov. Cotopaxi: Cotopaxi N. P., NNE slope of Volcán Cotopaxi, alt. 4200–4250 m, 00°39′ S, 78°25′ W, Palice 4604 (PRA).—Peru: Prov. Canchis: Sibinasallma, Murmurani sector, W of la Laguna Sibinacocha, PR 566a, 566b (LPB). Dept. Arequipa: Prov. Caylloma, Cañon del Colca canyon, between Soro and Llahuar villages, alt. 2100 m, 15°34′41″S, 72°01′01″W, AF 10135, 10139 & MK 6107, 6108 (KRAM, UGDA). Dept. Cuzco: prov. Urubamba, valley of Rio Piri, 22 km (road distance) c. NW of Ollantaytambo, alt. 3400 m, 13°06′S, 72°22′W, Santesson P86: 17 (S). Dept. Lima: Prov. Huarochiri, valley of Rio Santa Eulalia, NE of Carampoma, alt. c. 3700 m, 11°38′S, 76°27′W, Santesson P24: 5 & R. Moberg (S). Dept. Junin: Prov. Tarma, c. 10 km (road distance) NNE of Palca, alt. c. 2600 m, 11°18′S, 75°32′W, Santesson P12: 60 & Moberg (S, mixed in specimen of L. diffusa).
Lepraria yunnaniana (Hue) Zahlbr
Syn. Lepraria nigrocincta Diederich et al., Bibliotheca Lichenologica 64: 78 (1997).
Chemistry. Divaricatic acid.
Notes. For the characteristics of the species see Aptroot et al. (Reference Aptroot, Diederich, Sérusiaux and Sipman1997), Laundon (Reference Laundon2008) and Kukwa & Flakus (Reference Kukwa and Flakus2009).
Lepraria yunnaniana is known from numerous localities in Africa, Asia, Australia, Papua New Guinea, Central America (Costa Rica) and South America (Bolivia, Ecuador) (e.g. Aptroot et al. Reference Aptroot, Diederich, Sérusiaux and Sipman1997; Aptroot & Feijen Reference Aptroot and Feijen2002; Kukwa Reference Kukwa2006a; Elix 2007, Reference Elix2009; Nöske et al. Reference Nöske, Mandl and Sipman2007; Laundon Reference Laundon2008; Nelsen & Gargas Reference Nelsen and Gargas2008; Kukwa & Flakus Reference Kukwa and Flakus2009). Here it is reported for the first time from Colombia, from humid montane forest, where it grew on moribund bryophytes.
Specimen examined. Colombia: Depto. del Antioquia: Municipio de Medellín, c. 14·5 km SE de Medellín hacia Rionegro, c. 06°10′N, 75°30′W, elev. 2000 m, Sastre-De Jesús & Churchill 898 (B).
We are greatly indebted to Dr Z. Palice (Průhonice) for allowing us to publish some of his data and for kind assistance with literature, to Dr Stephan G. Beck and Mrs Rosa Isela Moneses Q. (Herbario Nacional de Bolivia, Instituto de Ecología, Universidad Mayor de San Andrés, La Paz) for generous cooperation, and to the curators of B, H, and S for the loan of specimens. A. Flakus warmly thanks Dr Harrie J. M. Sipman (Berlin) for making specimens available for this study, and for kind hospitality during his visit to B. Two anonymous reviewers are thanked for making several helpful comments. This research received support from the National Centre for Research and Development (NCBiR) in Poland under the LIDER Programme for the years 2010–2013 (no. 92/L-1/09), and from the SYNTHESYS Project (DE-TAF-4939), which is financed by European Community Research Infrastructure Action under the FP6 ‘Structuring the European Research Area’ Programme.