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A new fertile species of Menegazzia and notes on two sorediate species from the Neotropics
Lichenologist 34(6): 503–508 (2002) doi:10.1006/lich.2002.0423, available online at http://www.idealibrary.com on
A new fertile species of Menegazzia and notes on two sorediate species from the Neotropics Jarle W. BJERKE Abstract: Menegazzia neotropica sp. nov. is described from upper montane forest remnants and paramos in Ecuador, Colombia and Venezuela. It is characterized by short-pedicellate apothecia with ridged margins that partly obscure the discs, and by narrow, glossy lobes with brown-black margins. Menegazzia dissecta is recorded from South America for the first time and M. terebrata is recorded from Colombia and Ecuador for the first time. 2002 The British Lichen Society. Published by Elsevier Science Ltd. All rights reserved.
Key words: Andes, lichens, Menegazzia, new species, South America.
Introduction
Materials and Methods
Menegazzia A. Massal. is a species-rich lichen genus with 59 species known from the Southern Hemisphere (Filson 1996; Malcolm & Galloway 1997; Galloway & Quilhot 1999; Krog 2000; Bjerke & Elvebakk 2001; James et al. 2001). Most of these species are confined to southern temperate forests (Santesson 1942; Kantvilas & James 1987), and the number of species declines abruptly towards tropical and northern temperate biomes. Menegazzia terebrata, which was reported from the Venezuelan Andes by López Figueiras (1986) and included in the checklist from the same area (Marcano et al. 1996), is the only species known from South America north of the Tropic of Capricorn. Herbarium specimens of Menegazzia from the Neotropics were studied and compared with species from other areas. As a result of these studies a new species is described, and new records of two other species are reported.
Specimens of Menegazzia from tropical South America were obtained from the following herbaria: B, GB, MERF and S. Material collected by Nicole Nöske in Ecuador was duplicated, and the duplicates examined have been sent to B. Material for comparison was obtained from GZU, HO and UPS, and comparisons were also made with material from southern South America cited by Bjerke (2001) and Bjerke & Elvebakk (2001). Thin-layer chromatography of acetone extracts was performed using standardized procedures (Culberson 1972; White & James 1985).
J. W. Bjerke: Department of Biology, Faculty of Science, University of Tromsø, N-9037 Tromsø, Norway. 0024–2829/02/060503+06 $35.00/0
The Species Menegazzia dissecta (Rass.) Hafellner Fritschiana 12: 18 (1997).—M. pertusa f. dissecta Rass., Nov. Sist. Niz. Rast. 1: 247 (1964).—M. terebrata var. dissecta (Rass.) Poelt, Bestimmungsschl. eur. Flechten: 756 (1969).—M. pertusa var. dissecta (Rass.) Rass. Nov. Sist. Niz. Rast. 10: 198 (1973); type: Russia, Oriens extremus, Sichote-Alinj austr., publicum reservatum Sudzuchinskij, 29 August 1944, Zhudova & Pokrovskaja (LE—holotype).
(Fig. 1) Thallus corticolous, closely attached, forming irregular, radiating patches or rosettes to 9 cm wide. Lobes 0·8–2·5 mm wide, inflated, contiguous to margins, partly
2002 The British Lichen Society. Published by Elsevier Science Ltd. All rights reserved.
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contiguous; soredia granular, white. Lower surface black, pleated. Cavity white above, black below. Apothecia and pycnidia not seen. Chemistry. Atranorin (minor), stictic acid (major), menegazziaic acid (major to minor), cryptostictic acid (minor), and constictic acid (major).
F. 1. Menegazzia dissecta (R. Santesson, A. Tehler & G. Thor P105:34). A, sorediate pustules protruding from the lobe apices; B, detail of sorediate, lacerate pustules. Scales: A=1·5 mm; B=0·76 mm.
overlapping centrally, almost without intrathalline spaces, subdichotomously branched at apices, more irregularly branched centrally, slightly contorted. Upper surface slightly convex, grey, greyish green and glossy at apices, blackened centrally and along margins, emaculate or slightly maculate, epruinose, sorediate. Perforations scattered, often minute, 0·05–1·1 mm wide, becoming ellipsoid; edge of perforations flat or inrolled. Soralia up to 2·0 mm diam.; formed on laminal and marginal pustules that rupture at an early stage, up to 2·5 mm tall, lacerate or flanged, with an open duct to the cavity, often clustered; clusters up to 5 mm wide; soralia on parallel lobes often
Notes. Menegazzia dissecta shows little morphological and chemical variation. The specimen from Peru is slightly glossier and has slightly shorter pustules than the Austrian specimens seen, but gross morphology is similar. It is distinguished from M. terebrata by the lacerate soralia that are formed on pustules, the more irregularly branched lobes, and the slightly more blackened margins. Menegazzia dissecta shares a similar morphology of soralia to some Southern Hemisphere species. The Australasian species M. nothofagi (Zahlbr.) P. James & D. Galloway has isidia-like vesicles or pustules that finally become sorediate and ruptured (Galloway 1985). It is distinguished from M. dissecta by the smaller thalli and lobes, the numerous perforations and the slightly concave upper surface. The Tasmanian endemic M. kantvilasii P. James produces lacerate soralia on laminal pustules (James & Galloway 1992), but is distinguished from M. dissecta by having smaller lobes, perforations with elevated rims, two types of soralia (one associated with perforations), and fatty acids. Except for the fatty acids in M. kantvilasii, the three species M. nothofagi, M. kantvilasii and M. dissecta have identical chemistry. The Chilean species M. wandae Bjerke is distinguished from M. dissecta by having slightly shorter pustules, that are not clustered, and thamnolic acid as a major medullary constituent (Bjerke 2001). The New Guinean species M. efflorescens P. James, Aptroot, Sérus. & Diedrich seems to have a similar chemistry and morphology as M. dissecta (see description by James et al. 2001), but material of this species has not been seen by the present author.
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Menegazzia neotropica—Bjerke
Menegazzia dissecta was described as M. pertusa f. dissecta (Rassadina 1964), but was subsequently raised to varietal rank by both Poelt (1969) and Rassadina (1973). Poelt (1969) indicated that there was no continuum between the maniciform soralium type of M. terebrata and the lacerate soralium type of M. dissecta. Based on Austrian specimens, Hafellner (1997) reached the same conclusion, and raised the taxon to species rank. The comparisons with Southern Hemisphere taxa indicate that M. dissecta is more closely related to M. nothofagi and M. kantvilasii than to M. terebrata, and that the recognition of M. dissecta at species rank is well justified. Distribution and habitat ecology. The only known locality of M. dissecta in South America is in Peru, in an upper montane forest at 3000 m a.s.l. where it is associated with Parmotrema spp., Leptogium spp., Usnea spp., crustose lichens and several mosses. No specimens of Menegazzia were found during the author’s field studies in 2001 in the same region of Peru, at altitudes both below and above 3000 m, but not so close to the Amazon basin. Rassadina (1964) reported this taxon from the eastern part of Russia close to the Pacific Ocean, Japan, China, Central and Northwest Europe [southern Scandinavia (mapped locality in Sweden), the Alps, the Pyrenees, Hungary, former Czechoslovakia, Poland and Germany], USA (Alaska and Hawaii), Canada, Mexico and Madagascar. However, only one specimen, the type, was listed. Poelt (1969) reported it from the German part of the Alps. Hafellner (1997) reported it from Steiermark in Austria and both specimens listed were seen during the present study. A specimen from Austria not listed by Hafellner (1997) and a specimen from Japan are included below. Selected specimens examined. Peru: Depto. Cusco: Provincia de Paucartambo, road Paucartambo-Pillcopata, 15 km (road distance) NE of turn to Tres Cruces, 1981, R. Santesson, A. Tehler & G. Thor P105:34 (S).— Austria: Steiermark: Nördliche Kalkalpen, N of Toplitzsee, 1999, J. Hafellner 49505 (GZU 02-99).— Japan: Honshu: Mt. Fuji, 25 April 1964, G. Degelius s.n. (UPS).
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Menegazzia neotropica Bjerke sp. nov. Differt a Menegazzia fissicarpa lobis nitidis et concavis, marginibus loborum piceis, marginibus foraminum deorsum curvatis. Typus: Colombia, Depto. Cauca, Municipio Puracé, Cordillera Central, along road La Plata-Puracé, 2 km W of Laguna San Rafael, alt. 3300 m., on Diplostephium in c. 3 m tall, open scrub in paramo bog, 6 October 1984, J. Aguirre C. & H. J. M. Sipman 5983 (B 44745—holotypus).
(Fig. 2) Thallus corticolous, closely to loosely attached, forming irregular rosettes, often on small twigs or among mosses and then very irregular and fragmentary, to 8 cm diam. Lobes numerous, contiguous and partly imbricate especially towards centre, narrow, 0·6–1·5(–1·8) mm wide, not much inflated, with numerous, small, toe-like lateral lobes. Upper surface grey, brown-black towards centre and along margins, usually slightly concave, smooth, emaculate, epruinose, glossy both at apices and towards centre; apices slightly raised, pale to dark brown, rounded. Perforations numerous, plane to slightly inrolled, circular, 0·1–0·6(–1·0) mm wide. Soralia and isidia absent. Lower surface black, dark brown and glossy at apices, pleated. Cavity creamy white on both upper and lower sides, becoming black, tomentose. Apothecia sparse to abundant, laminal, often clustered, unevenly oval, rarely circular, 0·4–1·3 mm wide, usually short-pedicellate; pedicel and thalline exciple slightly inflated, becoming fluted with sinuous vertical ridges; exciplewavy, often becoming brownish, inrolled, partly obscuring the disc; disc glossy, concave, epruinose, red-brown. Epithecium red-brown, 20–28 m thick; thecium colourless, 80–150 m tall; hypothecium brown, 30–40 m thick, unchanged in K. Asci 2-spored at maturity, 108–14040– 78 m. Ascospores ellipsoid, 53–7430– 47 m, thick-walled; wall 4·5–7·5 m thick. Pycnidia sparse; ostiole black; conidia 5–70·5 m. Chemistry. Atranorin (major to minor), stictic acid (major), constictic acid (major),
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by concave, glossy lobes with brown-black margins, and by the small perforations with inrolled rims. It shows little in the way of morphological variation. Apothecia are rarely only slightly ridged, for example the specimen from Venezuela (López-Figueiras 30184) has only a few apothecia with the characteristic ridged margins, and most of the discs are visible. The Tasmanian endemic M. corrugata P. James and the Australian endemic M. fissicarpa P. James are also characterized by apothecia with uneven, ridged or corrugate margins (James & Galloway 1992), which resemble the apothecia of M. neotropica, and they also have a similar chemistry to M. neotropica. Menegazzia corrugata is distinguished from M. neotropica by more markedly inflated and much larger apothecia, by the broad, rugosereticulate, convex lobes, and by the conical perforations. Menegazzia fissicarpa differs from M. neotropica by the conical perforations, slightly wider and more convex lobes with primarily grey margins, and by the presence of an unidentified medullary substance that is UV+ white. The apothecia of the specimens examined of M. fissicarpa are generally larger (1·0–3·5 mm wide) than the size reported by James & Galloway (1992; 0·3–0·7 mm wide), and larger than those of M. neotropica. None of the southern South American species shares morphological and anatomical features with M. neotropica.
F. 2. Menegazzia neotropica (holotype). A, part of thallus; B, cluster of apothecia with margins rolling inwards and hiding the discs; C, detail of ridged pedicel of the apothecium. Scales: A=2·2 mm; B=0·56 mm; C=0·34 mm.
cryptostictic acid (minor) and menegazziaic acid (trace). Notes. Menegazzia neotropica is characterized by the short-pedicellate apothecia with prominent, sinuous, corrugate, ridged, inrolled exciples that partly obscure the discs,
Distribution and habitat ecology. Menegazzia neotropica is known from latitude 315 S in southern Ecuador to approximately latitude 840 N in western Venezuela. It is confined to high altitudes (between 3100 and 3800 m a.s.l.) along the Andes, where it occurs in forest and woodland remnants within the paramo ecosystems (see Luteyn 1992; Sipman 1992). It grows on trunks and twigs of smooth-barked trees and shrubs within the subparamo and grass paramo belts. It associates with a wide range of other lichens and mosses, for example Usnea spp., Sticta spp., Hypotrachyna spp., Parmotrema spp. and Oropogon spp.
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Menegazzia neotropica—Bjerke
Additional specimens examined. Colombia: Depto. Cundinamarca: Municipio Guasca, Páramo de Guasca, Pena Negra, 1984, J. Aguirre C. & H. J. M. Sipman 5097 (B 44747); Municipio Supatá, Alto El Tablazo, along track from radar station to Supatá, 1984, J. Aguirre C. & H. J. M. Sipman 5330 (B 44749). Depto. Risaralda: Municipio Santa Rosa de Cabal, Camino real between Termales de Santa Rosa and Hda. La Sierra, Finca Berlin, 1986, J. Wolf 993 (B 91826); Municipio Santa Rosa de Cabal, W-slope of Cordillera Central, c. 500 m S of Finca La Sierra, 1984, J. Aguirre C. & H. J. M. Sipman 5435 (B 44746).—Ecuador: Provincia de Azuay: Páramo de Tinajillas, 22 km S of Cumbre on road Cuenca-Loja, 1985, L. Arvidsson 6873, M. Lindqvist & M. Lindström (GB 34731).—Venezuela: Estado Mérida: Páramo de San José de Acequias, near Zanjón del Cupís, s. annum, M. López-Figueiras 30184 (B 48724).
Menegazzia terebrata (Hoffm.) A. Massal. Neagen. Lich.: 3 (1854); for synonyms, see Rassadina (1964) and Santesson (1993).
Notes. Menegazzia terebrata is characterized by a closely attached thallus rosette having radiating, contiguous lobes with marginal or laminal convex soralia that occasionally show an open duct to the cavity. Perforations are small, often less than 0·5 mm wide, and regularly distributed. Stictic, constictic, menegazziaic and cryptostictic acids are the major medullary constituents. Apothecia are not known from South American material. For a more detailed description, see for example Poelt (1969) or Purvis et al. (1992). Soralia appear to be rarer in the South American specimens than in northern temperate specimens. Menegazzia subpertusa P. James & D. J. Galloway is the southern temperate species that most closely resembles M. terebrata, but the latter has a smoother thallus surface, the morphology of the soralia is different (Galloway 1985), the ascospores are larger, and the perforations are smaller. Most older reports of M. terebrata (as M. pertusa) from Australia are referable to M. subpertusa (Filson 1996). Distribution and habitat ecology. It is a rare species in Ecuador, growing on trunks of trees in montane tropical rainforests in the south of the country, and has so far been collected only at altitudes between 2100 and
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2200 m a.s.l., which is below the paramo belt. The only specimen examined from Colombia is from the subparamo belt where it grows on shrubby plants and probably also on small trees. In Venezuela, it grows in montane forests and paramos, between 2300 and 3100 m a.s.l. (Marcano et al. 1996). To the best of my knowledge, it has not been reported from Ecuador and Colombia, previously. Specimens examined. Colombia: Depto. Cundinamarca: Municipio Guasca, Páramo de Guasca, Pena Negra, 1984, J. Aguirre C. & H. J. M. Sipman 5184b (B 44748).—Ecuador: Provincia de Zamora-Chinchipe: Reserva Biológica San Francisco, between Loja and Zamora, at the northern limit of Parque Nacional de Podocarpus, 2001, N. Nöske 231, 2071, 2072 (B).— Venezuela: Estado Tachira: Betania, upper part of Valle de Tamá, 1981, M. López-Figueiras 25966 (MERF). I am grateful to Dr Harrie Sipman, Freie Universität Berlin, for his efforts in bringing back the material housed in B that he and his colleagues collected in tropical South America, and Ms Nicole Nöske, University of Göttingen, for making duplicates of her sparse and valuable material available and for giving information on the abundance of Menegazzia in southern Ecuador. Dr Lars Arvidsson, University of Gothenburg, kindly gave advice on where to find the material he and his colleagues collected in Ecuador. I am indebted to the curators of B, GB, GZU, HO, MERF, S and UPS for placing material at my disposal, to Ms Gunvor Granaas, University of Tromsø, for giving advice on photography, to Ms Anna Johnsen, Tromsø and Arkhangelsk, for translation of Rassadina’s papers, and to Dr Arve Elvebakk, University of Tromsø, for comments on the manuscript and for suggesting the Latin name for the new species. R Bjerke, J. W. (2001) A new sorediate species of Menegazzia (Parmeliaceae, lichenized Ascomycota) from Chile. Lichenologist 33: 117–120. doi:10.1006/ lich.2000.0302. Bjerke, J. W. & Elvebakk, A. (2001) The sorediate species of the genus Menegazzia (Parmeliaceae, lichenized Ascomycotina) in southernmost South America. Mycotaxon 78: 363–392. Culberson, C. F. (1972) Improved conditions and new data for the identification of lichen products by a standardized thin layer chromatographic method. Journal of Chromatography 72: 113–125. Filson, R. B. (1996) Checklist of Australian lichens and allied fungi. Flora of Australia Supplementary Series 7: 1–204. Galloway, D. J. (1985) Flora of New Zealand. Lichens. Wellington: P. D. Hasselberg, Government Printer.
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Galloway, D. J. & Quilhot, W. (1999) [‘1998’] Checklist of Chilean lichen-forming and lichenicolous fungi. Gayana Botánica 55: 111–185. Hafellner, J. (1997) Materialien zur roten Liste gefährdeter Flechten Österreichs. Fritschiana 12: 1–32. James, P. W., Aptroot, A., Diedrich, P., Sipman, H. J. M. & Sérusiaux, E. (2001) New species of the lichen genus Menegazzia in New Guinea. Bibliotheca Lichenologica 78: 91–108. James, P. W. & Galloway, D. J. (1992) Menegazzia. Flora of Australia 54: 213–246. Kantvilas, G. & James, P. W. (1987) The macrolichens of Tasmanian rainforest: key and notes. Lichenologist 19: 1–28. Krog, H. (2000) Corticolous macrolichens of low montane rainforests and moist woodlands of eastern Tanzania. Sommerfeltia 28: 1–75. López-Figueiras, M. (1986) Censo de macrolíquenes de los estados Falcón, Lara, Mérida, Táchira, y Trujillo. Mérida: Facultad de Farmacia, Universidad de los Andes. Luteyn, J. L. (1992) Páramos: why study them? In Páramo: an Andean Ecosystem Under Human Influence (H. Balslev & J. L. Luteyn, eds): 1–14. London: Academic Press. Malcolm, W. M. & Galloway, D. J. (1997) New Zealand Lichens. Checklist, Key, and Glossary. Christchurch: Museum of New Zealand Te Papa Tongarewa, Caxton Press. Marcano, V., Méndez, A. M., Sipman, H. & Calderon, L. (1996) A first checklist of the lichen-forming
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fungi of the Venezuelan Andes. Tropical Bryology 12: 193–235. Poelt, J. (1969) Bestimmungsschlüssel europäischer Flechten. Lehre: Cramer. Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. W. & Moore, D. M. (eds) (1992) The Lichen Flora of Great Britain and Ireland. London: Natural History Museum Publications. Rassadina, K. A. (1964) Menegazzia Mass. in URSS. Novosti Sistematiki Nizsich rastenij [Novitates Systematicae Plantarum non Vascularium] 1: 235– 250. Rassadina, K. A. (1973) Formae et combinationes Parmeliacearum novae in determinatorio lichenum URSS, Fasc. 1 acceptae. Novosti Sistematiki Nizsich rastenij [Novitates Systematicae Plantarum non Vascularium] 10: 196–200. Santesson, R. (1942) The South American Menegazziae. Arkiv für Botanik 30A(11): 1–35. Santesson, R. (1993) The Lichens and Lichenicolous Fungi of Sweden and Norway. Lund: SBTförlaget. Sipman, H. (1992) The origin of the lichen flora of the Colombian páramos. In Páramo: an Andean Ecosystem Under Human Influence (H. Balslev & J. L. Luteyn, eds): 95–109. London: Academic Press. White, F. J. & James, P. W. (1985) A new guide to microchemical techniques for the identification of lichen substances. British Lichen Society Bulletin 57(suppl.): 1–41.
Accepted for publication 7 October 2002
A new fertile species of Menegazzia and notes on two sorediate species from the Neotropics Jarle W. BJERKE Abstract: Menegazzia neotropica sp. nov. is described from upper montane forest remnants and paramos in Ecuador, Colombia and Venezuela. It is characterized by short-pedicellate apothecia with ridged margins that partly obscure the discs, and by narrow, glossy lobes with brown-black margins. Menegazzia dissecta is recorded from South America for the first time and M. terebrata is recorded from Colombia and Ecuador for the first time. 2002 The British Lichen Society. Published by Elsevier Science Ltd. All rights reserved.
Key words: Andes, lichens, Menegazzia, new species, South America.
Introduction
Materials and Methods
Menegazzia A. Massal. is a species-rich lichen genus with 59 species known from the Southern Hemisphere (Filson 1996; Malcolm & Galloway 1997; Galloway & Quilhot 1999; Krog 2000; Bjerke & Elvebakk 2001; James et al. 2001). Most of these species are confined to southern temperate forests (Santesson 1942; Kantvilas & James 1987), and the number of species declines abruptly towards tropical and northern temperate biomes. Menegazzia terebrata, which was reported from the Venezuelan Andes by López Figueiras (1986) and included in the checklist from the same area (Marcano et al. 1996), is the only species known from South America north of the Tropic of Capricorn. Herbarium specimens of Menegazzia from the Neotropics were studied and compared with species from other areas. As a result of these studies a new species is described, and new records of two other species are reported.
Specimens of Menegazzia from tropical South America were obtained from the following herbaria: B, GB, MERF and S. Material collected by Nicole Nöske in Ecuador was duplicated, and the duplicates examined have been sent to B. Material for comparison was obtained from GZU, HO and UPS, and comparisons were also made with material from southern South America cited by Bjerke (2001) and Bjerke & Elvebakk (2001). Thin-layer chromatography of acetone extracts was performed using standardized procedures (Culberson 1972; White & James 1985).
J. W. Bjerke: Department of Biology, Faculty of Science, University of Tromsø, N-9037 Tromsø, Norway. 0024–2829/02/060503+06 $35.00/0
The Species Menegazzia dissecta (Rass.) Hafellner Fritschiana 12: 18 (1997).—M. pertusa f. dissecta Rass., Nov. Sist. Niz. Rast. 1: 247 (1964).—M. terebrata var. dissecta (Rass.) Poelt, Bestimmungsschl. eur. Flechten: 756 (1969).—M. pertusa var. dissecta (Rass.) Rass. Nov. Sist. Niz. Rast. 10: 198 (1973); type: Russia, Oriens extremus, Sichote-Alinj austr., publicum reservatum Sudzuchinskij, 29 August 1944, Zhudova & Pokrovskaja (LE—holotype).
(Fig. 1) Thallus corticolous, closely attached, forming irregular, radiating patches or rosettes to 9 cm wide. Lobes 0·8–2·5 mm wide, inflated, contiguous to margins, partly
2002 The British Lichen Society. Published by Elsevier Science Ltd. All rights reserved.
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contiguous; soredia granular, white. Lower surface black, pleated. Cavity white above, black below. Apothecia and pycnidia not seen. Chemistry. Atranorin (minor), stictic acid (major), menegazziaic acid (major to minor), cryptostictic acid (minor), and constictic acid (major).
F. 1. Menegazzia dissecta (R. Santesson, A. Tehler & G. Thor P105:34). A, sorediate pustules protruding from the lobe apices; B, detail of sorediate, lacerate pustules. Scales: A=1·5 mm; B=0·76 mm.
overlapping centrally, almost without intrathalline spaces, subdichotomously branched at apices, more irregularly branched centrally, slightly contorted. Upper surface slightly convex, grey, greyish green and glossy at apices, blackened centrally and along margins, emaculate or slightly maculate, epruinose, sorediate. Perforations scattered, often minute, 0·05–1·1 mm wide, becoming ellipsoid; edge of perforations flat or inrolled. Soralia up to 2·0 mm diam.; formed on laminal and marginal pustules that rupture at an early stage, up to 2·5 mm tall, lacerate or flanged, with an open duct to the cavity, often clustered; clusters up to 5 mm wide; soralia on parallel lobes often
Notes. Menegazzia dissecta shows little morphological and chemical variation. The specimen from Peru is slightly glossier and has slightly shorter pustules than the Austrian specimens seen, but gross morphology is similar. It is distinguished from M. terebrata by the lacerate soralia that are formed on pustules, the more irregularly branched lobes, and the slightly more blackened margins. Menegazzia dissecta shares a similar morphology of soralia to some Southern Hemisphere species. The Australasian species M. nothofagi (Zahlbr.) P. James & D. Galloway has isidia-like vesicles or pustules that finally become sorediate and ruptured (Galloway 1985). It is distinguished from M. dissecta by the smaller thalli and lobes, the numerous perforations and the slightly concave upper surface. The Tasmanian endemic M. kantvilasii P. James produces lacerate soralia on laminal pustules (James & Galloway 1992), but is distinguished from M. dissecta by having smaller lobes, perforations with elevated rims, two types of soralia (one associated with perforations), and fatty acids. Except for the fatty acids in M. kantvilasii, the three species M. nothofagi, M. kantvilasii and M. dissecta have identical chemistry. The Chilean species M. wandae Bjerke is distinguished from M. dissecta by having slightly shorter pustules, that are not clustered, and thamnolic acid as a major medullary constituent (Bjerke 2001). The New Guinean species M. efflorescens P. James, Aptroot, Sérus. & Diedrich seems to have a similar chemistry and morphology as M. dissecta (see description by James et al. 2001), but material of this species has not been seen by the present author.
2002
Menegazzia neotropica—Bjerke
Menegazzia dissecta was described as M. pertusa f. dissecta (Rassadina 1964), but was subsequently raised to varietal rank by both Poelt (1969) and Rassadina (1973). Poelt (1969) indicated that there was no continuum between the maniciform soralium type of M. terebrata and the lacerate soralium type of M. dissecta. Based on Austrian specimens, Hafellner (1997) reached the same conclusion, and raised the taxon to species rank. The comparisons with Southern Hemisphere taxa indicate that M. dissecta is more closely related to M. nothofagi and M. kantvilasii than to M. terebrata, and that the recognition of M. dissecta at species rank is well justified. Distribution and habitat ecology. The only known locality of M. dissecta in South America is in Peru, in an upper montane forest at 3000 m a.s.l. where it is associated with Parmotrema spp., Leptogium spp., Usnea spp., crustose lichens and several mosses. No specimens of Menegazzia were found during the author’s field studies in 2001 in the same region of Peru, at altitudes both below and above 3000 m, but not so close to the Amazon basin. Rassadina (1964) reported this taxon from the eastern part of Russia close to the Pacific Ocean, Japan, China, Central and Northwest Europe [southern Scandinavia (mapped locality in Sweden), the Alps, the Pyrenees, Hungary, former Czechoslovakia, Poland and Germany], USA (Alaska and Hawaii), Canada, Mexico and Madagascar. However, only one specimen, the type, was listed. Poelt (1969) reported it from the German part of the Alps. Hafellner (1997) reported it from Steiermark in Austria and both specimens listed were seen during the present study. A specimen from Austria not listed by Hafellner (1997) and a specimen from Japan are included below. Selected specimens examined. Peru: Depto. Cusco: Provincia de Paucartambo, road Paucartambo-Pillcopata, 15 km (road distance) NE of turn to Tres Cruces, 1981, R. Santesson, A. Tehler & G. Thor P105:34 (S).— Austria: Steiermark: Nördliche Kalkalpen, N of Toplitzsee, 1999, J. Hafellner 49505 (GZU 02-99).— Japan: Honshu: Mt. Fuji, 25 April 1964, G. Degelius s.n. (UPS).
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Menegazzia neotropica Bjerke sp. nov. Differt a Menegazzia fissicarpa lobis nitidis et concavis, marginibus loborum piceis, marginibus foraminum deorsum curvatis. Typus: Colombia, Depto. Cauca, Municipio Puracé, Cordillera Central, along road La Plata-Puracé, 2 km W of Laguna San Rafael, alt. 3300 m., on Diplostephium in c. 3 m tall, open scrub in paramo bog, 6 October 1984, J. Aguirre C. & H. J. M. Sipman 5983 (B 44745—holotypus).
(Fig. 2) Thallus corticolous, closely to loosely attached, forming irregular rosettes, often on small twigs or among mosses and then very irregular and fragmentary, to 8 cm diam. Lobes numerous, contiguous and partly imbricate especially towards centre, narrow, 0·6–1·5(–1·8) mm wide, not much inflated, with numerous, small, toe-like lateral lobes. Upper surface grey, brown-black towards centre and along margins, usually slightly concave, smooth, emaculate, epruinose, glossy both at apices and towards centre; apices slightly raised, pale to dark brown, rounded. Perforations numerous, plane to slightly inrolled, circular, 0·1–0·6(–1·0) mm wide. Soralia and isidia absent. Lower surface black, dark brown and glossy at apices, pleated. Cavity creamy white on both upper and lower sides, becoming black, tomentose. Apothecia sparse to abundant, laminal, often clustered, unevenly oval, rarely circular, 0·4–1·3 mm wide, usually short-pedicellate; pedicel and thalline exciple slightly inflated, becoming fluted with sinuous vertical ridges; exciplewavy, often becoming brownish, inrolled, partly obscuring the disc; disc glossy, concave, epruinose, red-brown. Epithecium red-brown, 20–28 m thick; thecium colourless, 80–150 m tall; hypothecium brown, 30–40 m thick, unchanged in K. Asci 2-spored at maturity, 108–14040– 78 m. Ascospores ellipsoid, 53–7430– 47 m, thick-walled; wall 4·5–7·5 m thick. Pycnidia sparse; ostiole black; conidia 5–70·5 m. Chemistry. Atranorin (major to minor), stictic acid (major), constictic acid (major),
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by concave, glossy lobes with brown-black margins, and by the small perforations with inrolled rims. It shows little in the way of morphological variation. Apothecia are rarely only slightly ridged, for example the specimen from Venezuela (López-Figueiras 30184) has only a few apothecia with the characteristic ridged margins, and most of the discs are visible. The Tasmanian endemic M. corrugata P. James and the Australian endemic M. fissicarpa P. James are also characterized by apothecia with uneven, ridged or corrugate margins (James & Galloway 1992), which resemble the apothecia of M. neotropica, and they also have a similar chemistry to M. neotropica. Menegazzia corrugata is distinguished from M. neotropica by more markedly inflated and much larger apothecia, by the broad, rugosereticulate, convex lobes, and by the conical perforations. Menegazzia fissicarpa differs from M. neotropica by the conical perforations, slightly wider and more convex lobes with primarily grey margins, and by the presence of an unidentified medullary substance that is UV+ white. The apothecia of the specimens examined of M. fissicarpa are generally larger (1·0–3·5 mm wide) than the size reported by James & Galloway (1992; 0·3–0·7 mm wide), and larger than those of M. neotropica. None of the southern South American species shares morphological and anatomical features with M. neotropica.
F. 2. Menegazzia neotropica (holotype). A, part of thallus; B, cluster of apothecia with margins rolling inwards and hiding the discs; C, detail of ridged pedicel of the apothecium. Scales: A=2·2 mm; B=0·56 mm; C=0·34 mm.
cryptostictic acid (minor) and menegazziaic acid (trace). Notes. Menegazzia neotropica is characterized by the short-pedicellate apothecia with prominent, sinuous, corrugate, ridged, inrolled exciples that partly obscure the discs,
Distribution and habitat ecology. Menegazzia neotropica is known from latitude 315 S in southern Ecuador to approximately latitude 840 N in western Venezuela. It is confined to high altitudes (between 3100 and 3800 m a.s.l.) along the Andes, where it occurs in forest and woodland remnants within the paramo ecosystems (see Luteyn 1992; Sipman 1992). It grows on trunks and twigs of smooth-barked trees and shrubs within the subparamo and grass paramo belts. It associates with a wide range of other lichens and mosses, for example Usnea spp., Sticta spp., Hypotrachyna spp., Parmotrema spp. and Oropogon spp.
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Menegazzia neotropica—Bjerke
Additional specimens examined. Colombia: Depto. Cundinamarca: Municipio Guasca, Páramo de Guasca, Pena Negra, 1984, J. Aguirre C. & H. J. M. Sipman 5097 (B 44747); Municipio Supatá, Alto El Tablazo, along track from radar station to Supatá, 1984, J. Aguirre C. & H. J. M. Sipman 5330 (B 44749). Depto. Risaralda: Municipio Santa Rosa de Cabal, Camino real between Termales de Santa Rosa and Hda. La Sierra, Finca Berlin, 1986, J. Wolf 993 (B 91826); Municipio Santa Rosa de Cabal, W-slope of Cordillera Central, c. 500 m S of Finca La Sierra, 1984, J. Aguirre C. & H. J. M. Sipman 5435 (B 44746).—Ecuador: Provincia de Azuay: Páramo de Tinajillas, 22 km S of Cumbre on road Cuenca-Loja, 1985, L. Arvidsson 6873, M. Lindqvist & M. Lindström (GB 34731).—Venezuela: Estado Mérida: Páramo de San José de Acequias, near Zanjón del Cupís, s. annum, M. López-Figueiras 30184 (B 48724).
Menegazzia terebrata (Hoffm.) A. Massal. Neagen. Lich.: 3 (1854); for synonyms, see Rassadina (1964) and Santesson (1993).
Notes. Menegazzia terebrata is characterized by a closely attached thallus rosette having radiating, contiguous lobes with marginal or laminal convex soralia that occasionally show an open duct to the cavity. Perforations are small, often less than 0·5 mm wide, and regularly distributed. Stictic, constictic, menegazziaic and cryptostictic acids are the major medullary constituents. Apothecia are not known from South American material. For a more detailed description, see for example Poelt (1969) or Purvis et al. (1992). Soralia appear to be rarer in the South American specimens than in northern temperate specimens. Menegazzia subpertusa P. James & D. J. Galloway is the southern temperate species that most closely resembles M. terebrata, but the latter has a smoother thallus surface, the morphology of the soralia is different (Galloway 1985), the ascospores are larger, and the perforations are smaller. Most older reports of M. terebrata (as M. pertusa) from Australia are referable to M. subpertusa (Filson 1996). Distribution and habitat ecology. It is a rare species in Ecuador, growing on trunks of trees in montane tropical rainforests in the south of the country, and has so far been collected only at altitudes between 2100 and
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2200 m a.s.l., which is below the paramo belt. The only specimen examined from Colombia is from the subparamo belt where it grows on shrubby plants and probably also on small trees. In Venezuela, it grows in montane forests and paramos, between 2300 and 3100 m a.s.l. (Marcano et al. 1996). To the best of my knowledge, it has not been reported from Ecuador and Colombia, previously. Specimens examined. Colombia: Depto. Cundinamarca: Municipio Guasca, Páramo de Guasca, Pena Negra, 1984, J. Aguirre C. & H. J. M. Sipman 5184b (B 44748).—Ecuador: Provincia de Zamora-Chinchipe: Reserva Biológica San Francisco, between Loja and Zamora, at the northern limit of Parque Nacional de Podocarpus, 2001, N. Nöske 231, 2071, 2072 (B).— Venezuela: Estado Tachira: Betania, upper part of Valle de Tamá, 1981, M. López-Figueiras 25966 (MERF). I am grateful to Dr Harrie Sipman, Freie Universität Berlin, for his efforts in bringing back the material housed in B that he and his colleagues collected in tropical South America, and Ms Nicole Nöske, University of Göttingen, for making duplicates of her sparse and valuable material available and for giving information on the abundance of Menegazzia in southern Ecuador. Dr Lars Arvidsson, University of Gothenburg, kindly gave advice on where to find the material he and his colleagues collected in Ecuador. I am indebted to the curators of B, GB, GZU, HO, MERF, S and UPS for placing material at my disposal, to Ms Gunvor Granaas, University of Tromsø, for giving advice on photography, to Ms Anna Johnsen, Tromsø and Arkhangelsk, for translation of Rassadina’s papers, and to Dr Arve Elvebakk, University of Tromsø, for comments on the manuscript and for suggesting the Latin name for the new species. R Bjerke, J. W. (2001) A new sorediate species of Menegazzia (Parmeliaceae, lichenized Ascomycota) from Chile. Lichenologist 33: 117–120. doi:10.1006/ lich.2000.0302. Bjerke, J. W. & Elvebakk, A. (2001) The sorediate species of the genus Menegazzia (Parmeliaceae, lichenized Ascomycotina) in southernmost South America. Mycotaxon 78: 363–392. Culberson, C. F. (1972) Improved conditions and new data for the identification of lichen products by a standardized thin layer chromatographic method. Journal of Chromatography 72: 113–125. Filson, R. B. (1996) Checklist of Australian lichens and allied fungi. Flora of Australia Supplementary Series 7: 1–204. Galloway, D. J. (1985) Flora of New Zealand. Lichens. Wellington: P. D. Hasselberg, Government Printer.
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Galloway, D. J. & Quilhot, W. (1999) [‘1998’] Checklist of Chilean lichen-forming and lichenicolous fungi. Gayana Botánica 55: 111–185. Hafellner, J. (1997) Materialien zur roten Liste gefährdeter Flechten Österreichs. Fritschiana 12: 1–32. James, P. W., Aptroot, A., Diedrich, P., Sipman, H. J. M. & Sérusiaux, E. (2001) New species of the lichen genus Menegazzia in New Guinea. Bibliotheca Lichenologica 78: 91–108. James, P. W. & Galloway, D. J. (1992) Menegazzia. Flora of Australia 54: 213–246. Kantvilas, G. & James, P. W. (1987) The macrolichens of Tasmanian rainforest: key and notes. Lichenologist 19: 1–28. Krog, H. (2000) Corticolous macrolichens of low montane rainforests and moist woodlands of eastern Tanzania. Sommerfeltia 28: 1–75. López-Figueiras, M. (1986) Censo de macrolíquenes de los estados Falcón, Lara, Mérida, Táchira, y Trujillo. Mérida: Facultad de Farmacia, Universidad de los Andes. Luteyn, J. L. (1992) Páramos: why study them? In Páramo: an Andean Ecosystem Under Human Influence (H. Balslev & J. L. Luteyn, eds): 1–14. London: Academic Press. Malcolm, W. M. & Galloway, D. J. (1997) New Zealand Lichens. Checklist, Key, and Glossary. Christchurch: Museum of New Zealand Te Papa Tongarewa, Caxton Press. Marcano, V., Méndez, A. M., Sipman, H. & Calderon, L. (1996) A first checklist of the lichen-forming
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fungi of the Venezuelan Andes. Tropical Bryology 12: 193–235. Poelt, J. (1969) Bestimmungsschlüssel europäischer Flechten. Lehre: Cramer. Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. W. & Moore, D. M. (eds) (1992) The Lichen Flora of Great Britain and Ireland. London: Natural History Museum Publications. Rassadina, K. A. (1964) Menegazzia Mass. in URSS. Novosti Sistematiki Nizsich rastenij [Novitates Systematicae Plantarum non Vascularium] 1: 235– 250. Rassadina, K. A. (1973) Formae et combinationes Parmeliacearum novae in determinatorio lichenum URSS, Fasc. 1 acceptae. Novosti Sistematiki Nizsich rastenij [Novitates Systematicae Plantarum non Vascularium] 10: 196–200. Santesson, R. (1942) The South American Menegazziae. Arkiv für Botanik 30A(11): 1–35. Santesson, R. (1993) The Lichens and Lichenicolous Fungi of Sweden and Norway. Lund: SBTförlaget. Sipman, H. (1992) The origin of the lichen flora of the Colombian páramos. In Páramo: an Andean Ecosystem Under Human Influence (H. Balslev & J. L. Luteyn, eds): 95–109. London: Academic Press. White, F. J. & James, P. W. (1985) A new guide to microchemical techniques for the identification of lichen substances. British Lichen Society Bulletin 57(suppl.): 1–41.
Accepted for publication 7 October 2002