CBibliotheca Lichenologica 108, p

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1 CBibliotheca Lichenologica 108, p Stuttgart, May 2012 Collema paramense, sp. nova, with notes on Collema in the High Andes Per M. JØRGENSEN 1 & Zdenĕk PALICE 2, 3 1 Botanical Museum, University of Bergen, Allégaten 41, 5007 Bergen, Norway 2 Institute of Botany, Czech Academy of Sciences, Průhonice, Czech Republic 3 Department of Botany, Faculty of Natural Sciences, Charles University, Benátská 2, Prague, Czech Republic; zdenek.palice@ibot.cas.cz Abstract: A terricolous species in the lichen genus Collema is described from c m altitude in the paramos of northern Ecuador. It appears to belong in the north-western Andean element and to be a vicariant of the arctic-alpine Collema ceraniscum which has a similar ecology. The rarity of the genus in the Andes is pointed out, for which there is no obvious reason. However, some new records from Ecuador are added: C. fasciculare (found at c m, its highest recorded locality) and C. tenax s.lat.; also C. limosum from the outskirts of Quito, new to South America. The commonest species in the region is apparently C. glaucophthalmum, reaching 4000 m, but mainly found at lower altitudes in the selvas nubladas. Keywords: Collema, Ecuador, rarity, paramo. Introduction The paramo is a special and most characteristic vegetation found at high altitude below the permanent snowline and above the forest-line (c m) in the northern Andes (see e.g. CLEEF 1978, CUATRECASAS 1968, LUTEYN 1999, VARESCHI 1970) and reasonably lichen-rich according to studies in Colombia and Venezuela (SIPMAN 1992, 1999). PALICE and co-workers have recently collected lichens in the less well-known paramos of Ecuador (ARVIDSSON 1991), which proved to contain several new species (JØRGENSEN & PALICE 2010), including a characteristic small gelatinous species of Collema which does not agree with any species described in the comprehensive monograph of DEGELIUS (1954, 1974) with a subsequent addition (DEGELIUS 1994), and will be described below. Furthermore there are very few records of Collema in the High Andes (see SIPMAN 1992, 1999), therefore some additional finds are worth noting. However, this does not alter the impression that the genus is surprisingly rare in the region, especially when compared with the closely related genus Leptogium (JØRGENSEN 1997). DEGELIUS (1974) only lists 19 species from South America, mostly from Brazil and the southernmost parts of the continent, of which four are endemic. Collema scabrum Degel., a species described by him later (DEGELIUS 1994), is the only Collema species so far known to occur exclusively in Ecuador. Twenty is the lowest species-number on world scale, except for that of the Arctic regions. He gives no explanation for this, nor can we J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, Germany eschweizerbart_xxx /2012/ $ 1.50

2 136 Per M. Jørgensen & Zdenĕk Palice Material and methods The new Ecuadorean material was collected by Z. PALICE and co-workers during recent expeditions and is deposited in PRA with duplicates where possible in QCA. It has been studied by standard microscopic methods and TLC. The new species Collema paramense P.M. Jørg. & Palice species nova (Fig.1) Collemae ceranisco similis, sed thallo tenui, crustaceo-squamuloso, nitido; asci octospori sporis hyalinis, majoris, non-cuboideis. TYPUS: ECUADOR, Carchi, volcan Chiles, wet paramo near road, c. 1.5 km SW of the top of the volcano, alt m, 13.VII.1999, Z. Palice 2796 (PRA, holotype; BG & QCA, isotypes). Thallus rather small, up to 2 cm diam, subcrustaceous to small-squamulose, applanate with shiny black surface, often covered in apothecia. Thallus in section μm thick with pseudocortex to 10 μm thick and dark brown gelatinous pigmentation externally. It is filled with short chains of Nostoc, individual cells 3 4 μm diam., surrounded by a loose weft of anastamosing hyphae, 2 3 μm wide. Apothecia numerous, up to 2 mm diam., initially flat and sessile, finally some what stalked and convex with brown disc and distinct exciple, μm broad, the paraplectenchymatous proper exciple often visible as a paler, internal ring from above. Subhymenium of intricating hyphae, often indistinct. Hymenium μm high, I+ constantly blue, of usually simple paraphyses with clavate, brown-pigmented apices. Asci subcylindrical, μm long, thickened apically with internal amyloid plug, 8-spored. Ascospores hyaline, muriform, ellipsoid with acute ends, μm (Fig. 2). Conidiomata not observed. Chemistry: No lichen acids found by TLC. Etymology: The Latin epithet derives from paramo, the Indian name for the habitat Habitat & distribution: This terricolous species, growing among mosses, appears to be restricted to wet paramos in the High Andes. It is as yet only known from two collections in northern Ecuador where it was collected in a drainage ditch among mosses in a rather acid paramo with Espeletia pycnophylla Cuatr. and in an open boggy paramo on decaying Azorella cushions. However, it may prove to have a wider distribution as this kind of habitat is found throughout the northern Andes, and it may belong in the north-western element (JØRGENSEN 1983); furthermore, it is easily overlooked. Taxonomic notes: The generic position of this new species is rather difficult to decide since the thallus has a poorly developed cortex which on closer inspection proved to be a pseudocortex. We have accordingly placed it in Collema rather than Leptogium, although we are aware of the artificial nature of today s separation of these genera (e.g. WEDIN et al. 2009) which appear to be in need of redefinition. As pointed out above, Collema species are surprisingly rare in the Andes (SIPMAN 1992, 1999), particularly at this altitude. Only one species, the very

3 Collema paramense, sp. nova, with notes on Collema in the High Andes 137 Fig. 1. Collema paramense, part of the type collection (PRA). Scale: 2 mm. different C. scabrum, is recorded from about this altitude, namely 3600 m in a pine plantation at the Cotopaxi volcano (DEGELIUS 1994); the corticolous C. glaucophthalmum Nyl. is also known to reach this high (see below). However, NYLANDER (1863) described a species, C. coccophylloides, from Colombia at 2000 m which DEGELIUS (1974) renamed C. calliobotrys var. coccophylliza (Zahlbr.) Degel. and placed in the C. occultatum group. We believe that our new species also belongs in this group, even if its spores are not cuboid as in the other species. It has considerable superficial resemblance to the arctic-alpine C. ceraniscum Nyl. which is also terricolous and not known from South America. (Its nearest localities we have located are in the Rockies: Montana, DEBOLT & MCCUNE 1993, and Colorado, ST. CLAIR et al ) This species is normally subfruticulose, but the growth-form described as C. arcticum Lynge (LYNGE 1926) has a more squamulose Fig. 2. Ascospore of Collema paramense. thallus which is not as shiny as that of C. paramense, and the spores differ in being smaller, cuboid and pale brownish, and are normally found in 4-spored asci. Additional material studied: Ecuador, Pichincha, Guamani (= paramo de la Virgen), moist paramo N of road (Quito) Pifo-Papallacta, c. 25 km WSW of Quito, alt m, 19.VI. 1999, Z. Palice 2273 (PRA).

4 138 Per M. Jørgensen & Zdenĕk Palice New records (1) Collema fasciculare (L.) F.H. Wigg. According to DEGELIUS (1974), this is one of the most widespread species of the genus, although there is only one previous record in this part of the Andes, a collection from the subalpine region collected in Bolivia by Mandon in The present specimen was collected on twigs of Chuquiragua jussieui Gmel. in a transitional type of paramo, a shrub and cushion paramo verging on a desert paramo. The vegetation was very sparse with distantly scattered caulescent plants and occasional shrubs on an almost bare, unstable volcanic substratum. In the very same locality, the endemic Ecuadorean lichen, Cetrariastrum ecuadoriense (R.Sant.) Sipman, has once been collected and is distributed in an exsiccate (VĔZDA 2003). The altitude is just above 4300 m, higher than any other Collema sp. recorded (cf. DEGELIUS 1974, who gives 3700 m as the maximum, interestingly for C. fasciculare, in Kenya). The occurrence of C. fasciculare at the Ecuador site was frequent and the specimens were well-developed with quite typical spores. Specimen studied: Ecuador, Cotopaxi, NNE slope of Volcan Cotopaxi, above the valley of quebrada Tañiloma, alt m, 28.VI.1999, Z. Palice 2911 (PRA). (2) Collema glaucophthalmum Nyl. This appears to be the commonest and most widespread species in this region, previously being recorded from several countries (Colombia, Ecuador, Peru, Bolivia, Chile, Argentina and Brazil, as well as Paraguay, with the type from Mexico); it is also known from the mountains of East Africa and Australasia. The var. glaucophthalmum is known from c m (in the selvas nubladas ) upwards to 4000 m (in Peru). The newly collected material grew as usual on bushes at m alt. These collections confirm the previous impression of a fairly common species in this region. Specimens studied: Ecuador, Chimborazo, Sangay, Hda. Guargualla-Hda. San Eduardo, valley of Quebrada Yulumpala, alt m, 19.VII.1999, Z. Palice 2452, 2526 (PRA); Pichincha, c. 25 km N of Quito, S-side of Pululagua caldera, along a descending trail just below the arris of the caldera, alt m, 15.VI.1999, Z. Palice 3451 & Z. Soldán (PRA). (3) Collema tenax (Sw.) Ach. (s. lat.) This species has never been reliably recorded from the Andes, and our specimen is unfortunately without apothecia which are so important for accurate identification. There is, however, no doubt that it belongs in this complex, since it has mostly marginal conidiomata just as in that species. The pycnoconidia are bacilliform 4 5 x m. The rather poorly developed small-squamulose specimen grew on soil in a lava flow at m. The second author, however, collected a specimen of this group with welldeveloped apothecia on trampled ground in the periphery of Quito, not situated

5 Collema paramense, sp. nova, with notes on Collema in the High Andes 139 in the high-andean zone. It differs clearly from the former in the nearly crustose, strongly blue-green (when collected) thallus with innate apothecia, indicative of C. limosum (Ach.) Ach., a fact confirmed by the 4-spored asci containing large, x 9 13 m, sparingly muriform to septate ascospores. This species is new to South America, though previously recorded from Mexico. Specimens studied: (of Collema tenax) Ecuador, Cotopaxi, NNE slope of volcano Cotopaxi, small hill situated in lava flow, alt m, 26.IV.1999, Z. Palice 2980 (PRA); (of Collema limosum) Ecuador, Pichincha, Quito- Cochacamba, W slope of Mt Rucu Pichincha, the periphery of Quito, alt m, 18.IV.1999, Z. Palice 3778 (PRA, QCA). Acknowledgements Z. Palice acknowledges financial support from a long-term research development project no. RVO and the Ministry of Education, Youth and Sports of the Czech Republic (no ), and thanks colleagues who participated in the Ecuadorean field trips. Both authors are indebted to Jan Berge and Beate Helle of Bergen for their assistance with the illustrations. References ARVIDSSON, L. (1991): Lichenological studies in Ecuador. In: GALLOWAY, D. J. (ed.). Tropical lichens, their systematics, conservation and ecology. Systematic Association, London, pp CUATRECASAS, J. (1968): Paramo vegetation and its life forms. Collegium Geographicum 9: CLEEF, A. M. (1978): Characteristics of neotropical paramo vegetation and its subantarctic relations. Erdwissenschaftliche Forschung 11: DEBOLT, A. & MCCUNE, B. (1993): Lichens of Glacier National Park, Montana. Bryologist 96: DEGELIUS, G. (1954): The lichen genus Collema in Europe. Symbolae Botanicae Upsalienses 13 (2): DEGELIUS, G. (1974): The lichen genus Collema. The extra-european species. Symbolae Botanicae Upsalienses 20 (2): DEGELIUS, G. (1994): Studies in the lichen family Collemataceae VII. Two new tropical Collema species. Nordic Journal of Botany 14: JØRGENSEN, P. M. (1983): Distribution patterns of lichens in the Pacific region. Australian Journal of Botany, Supplement Series 10: JØRGENSEN, P. M. (1997): Further notes on hairy Leptogium species. Symbolae Botanicae Upsalienses 32 (1): JØRGENSEN, P. M. & PALICE, Z. (2010): Additions to the lichen family Pannariaceae in Ecuador. Nordic Journal of Botany 28: LUTEYN, J. L. (ed.) (1999): Paramos, a checklist of plant diversity, geographical distribution and botanical literature. New York Botanical Garden, Bronx, 278 pp. LYNGE, B. (1926): Lichens from Bear Island (Bjørnøya) collected by Norwegian and Swedish expeditions, chiefly by Th. M. Fries during the Swedish Polar Expedition of Norske Videnskaps Akademis Skrifter. Resultater av de norske Statsunderstøttede Spitsbergenekspeditioner 1 (9): NYLANDER, W. (1863): Lichenographiae Novo-Granatensis Prodromus. Acta Societatis Scientarum Fennicae 7:

6 140 Per M. Jørgensen & Zdenĕk Palice SIPMAN, H. J. M. (1992): The origin of the lichen flora of the Colombian paramos. In: BALSLEV, H. & LUTEYN, J. L. (eds). An Andean ecosystem under human influence. New York Botanical Garden, Bronx, pp SIPMAN, H. J. M. (1999): Lichenes. In: LUTEYN, J. L. (ed.). Paramos, a checklist of plant diversity, geographical distribution and botanical literature. New York Botanical Garden, Bronx, pp ST. CLAIR, S. B., COOPER, L. S., STEWART, L. J., NEWBERRY, C. C. & ST. CLAIR, L. L. (1998): The lichen flora of Stony Pass, an alpine tundra sites in southwestern Colorado. In: GLENN, M. G., HARRIS, R. C., DIRIG, R. & COLE, M. S. (eds). Lichenographia Thomsoniana: North American lichenology in honor of John W. Thomson. Mycotaxon Ltd., Ithaca, pp VARESCHI, V. (1970): Flora de los paramos de Venezuela. Merida, 429 pp. VĔZDA, A. (2003): Lichenes rariores exsiccati. Fasc. 49 (numeris ). Brno, 4 pp. WEDIN, M., WIKLUND, E., JØRGENSEN, P. M. & EKMAN, S. (2009): Slippery when wet: Phylogeny and character evolution in the gelatinous cyanobacterial lichens (Peltigerales, Ascomycetes). Molecular Phylogenetics and Evolution 53: