A New Species of Megalaria from I|Les Kerguelen in the Southern

The Lichenologist 39(4): 371–375 (2007) 2007 British Lichen Society doi:10.1017/S0024282907007025 Printed in the United Kingdom

A new species of Megalaria from| Iles Kerguelen in the southern Indian Ocean

Alan M. FRYDAY

Abstract: The new species Megalaria allantoidea Fryday is described from the subantarctic| Iles Kerguelen in the southern Indian Ocean, where it grows over terricolous bryophytes or Azorella selago cushions. It differs from all other described species of Megalaria in having large, sausage-shaped ascospores. The systematics of the genus Megalaria is reviewed, ascus structure of some previously described species revised, and the position of the new species in the genus discussed. Key words: lichenized fungi, Ramalinaceae, subantarctic

Introduction |Iles Kerguelen is a group of remote islands in the southern Indian Ocean (Fig. 1A). They consist of a main island, roughly 120 km by 140 km and occupying 6675 km2, sur- rounded by approximately 300 other smaller islands, reefs and rocks, forming an archipelago of 7215 km2 and with elevations ranging from sea-level to 1850 m (Fig. 1B). Situated almost 4000 km from Africa and Australia and over 2000 km from Antarctica (the nearest land mass of any size),| Iles Kerguelen are subject to one of the most oceanic climates anywhere in the world, i.e. a low temperature range and high precipitation. The Kerguelen climate is characterized by a mean annual temperature of 4·6(C, varying between a monthly average of 2·3(C in winter and 7·9(C in summer, with rain or snow on most days, and almost constant winds. However, precipitation is rarely heavy, annual precipitation averaging only about 850 mm but, as winds blow continu- ously from the west, this is usually in the form of driving drizzle. Winds of 150 kmph are common, and gusting up to 200 kmph

F. 1. A, map of the southern section of the eastern Alan M. Fryday: Herbarium, Department of Plant Southern Hemisphere showing the position of| Iles Biology, Michigan State University, East Lansing, MI Kerguelen; B, map of| Iles Kerguelen showing known 48824-1312, USA. Email: [email protected] localities of Megalaria allantoidea.

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has been known (Varney 1926). Geologi- KOH (K) and lactophenol cotton-blue. The ascus cally, the islands are volcanic in origin and structure was studied in 0·15% aqueous IKI, both with- out prior treatment and after pretreatment with 10% composed almost entirely of flood basalt. A KOH. Measurements of ascospores and paraphyses full description of the islands and their biota were made in 10% KOH. is given by Mawson (1934). Nomenclature for apothecial pigments follows Meyer In the austral summer of 1971, Henry & Printzen (2000). Imshaug led an expedition to investigate the terrestrial flora of the islands. Vascular The Species plants were studied by Dr Stephen Young (Ohio, USA), bryophytes by John Engel Megalaria allantoidea Fryday sp. nov. (MSU), and lichens by Imshaug, Richard Harris (both MSU), and Geoffrey Bratt MycoBank No.: MB 510688 (Tasmania) (Imshaug 1971). Engel and A Megalaris ceteris ascosporis allantoideus, grandiori- Harris were at that time Imshaug’s graduate bus (40–)45(–50)7–8(–10) m differt; thallus students. Between them Imshaug, Harris bryophilus. Typus: Kerguelen,| Ile Longue, summit of le Kiosque, and Bratt made over 2500 collections from [c. 49(30.75#S69(53.0#E], 199 m, 18 March 1971, the SE section of the islands, representing by H. A. Imshaug 49161 (MSC—holotypus). far the largest collection of lichens ever made from the islands. These collections were (Figs2&3) returned to the herbarium of Michigan State University (MSC), where most received only Thallus effuse, appearing as a pale grey a preliminary examination before being crust that is actually necrotic, de-pigmented placed in storage in cardboard boxes. How- bryophytes, small white granules, 0·2– ever, recent awards from the US National 0·25 mm, occasionally present. Photobiont Science Foundation (NSF) have led to the chlorococcoid, cells globose, 10–15 m re-activation of this extensive collection diam. (Fryday & Prather 2001) and resulted in the Apothecia black, lecideine, orbicular, label data of the entire accessioned lichen adnate, flat even when old, 0·7–1·2 mm collection at MSC being entered into a data- diam., proper margin persistent, barely base and made available on-line (Johnson raised above the level of the disc, 0·05 mm et al. 2005). wide. Excipulum composed of radiating In 1998, Dr U. Søchting and his graduate hyphae with enlarged cells (10–15 m wide), student, R. Poulsen, also visited| Iles 140 m thick, inner 50 m dark blue Kerguelen to collect lichens. Among the (K+ brighter; HCl+ blue, N+ magenta; many collections from both expeditions that Cinereorufa-green), outer with dark red represented apparently undescribed taxa (K+ brighter, HCl+ brown, N+ orange- (Fryday 2004a & unpublished; Poulsen et al. brown; Atra-red) pigment on outer walls of 2001; Jørgensen 2001) were specimens of the hyphae, continuous under hypothecium. a bryophilous species that was clearly con- Hymenium hyaline, 120–150 m, streaked generic with the recently described Meg- with dark red (Atra-red) pigment; epi- alaria imshaugii Fryday and M. macrospora hymenial zone dark red-purple (Atra-red), Fryday, but with large sausage-shaped as- 10–15 m thick; subhymenium dark red cospores, and which is here described as new (Atra-red) above with thin dark blue-green to science. (Cinereorufa-green) zone below, 35–40 m. Paraphyses simple, readily separating, 2·0– 2·5 m thick, apices swollen to 5·0–6·5 m Materials and Methods with dark red (Atra-red) pigmented cap. Hypothecium 70 m thick, hyaline to pale Apothecial characteristics were examined by light microscopy on hand-cut sections mounted in water, blue-green, composed of periclinally arranged hyphae with large yellow oil inclu- 10% KOH (K), 50% HNO3 (N) or 0·15% aqueous IKI. Thallus sections were investigated in water, 10% sions. Asci cylindrical, 80–8517–20

F.2. Megalaria allantoidea, thallus and apothecia. Scale=1·0 mm.

(–40) m, Bacidia-type. Ascospores hyaline, 1-septate, allantoid, (40–)45(–50)7–8 (–10) m, (6–)8/ascus. Conidiomata not observed.

Chemistry.K,C,Pd,UV; thallus too scanty to be tested by TLC.

Etymology. The epithet is derived from ‘‘allant’’, Greek , o, a sausage like dish, and ‘‘–oides’’ Greek , like, resemble, and refers to the distinctive sausage-shaped ascospores of the new species.

Distribution and ecology. The new species is known only from the subantarctic| Iles Kerguelen, where it has been collected from several localities between 100–700 m above sea-level (Fig. 1B). All collections are from terricolous bryophytes or Azorella selago Hook. f. cushions. Associated lichens include species of Pannariaceae.

Systematic position. The genus Megalaria F.3. Megalaria allantoidea, ascospore. Scale=10 m. Hafellner (1984) was erected for the single

corticolous species M. grossa (Pers. ex Nyl.) Species of Megalaria typically have Hafellner and placed in the monogeneric ascospores less than 25 m in length with a family Megalariaceae Hafellner, to which length/breadth ratio of c. 2:1. Megalaria Kantvilas et al. (1999) later added a second grossa and M. macrospora have larger genus, Tasmidella Kantvilas, Hafellner & ascospores (up to 30 m and 36 m long, Elix. Megalaria was expanded by the respectively), but these are still not as long as addition of several other temperate corti- those of M. allantoidea and their length/ colous species, and Ekman & Tønsberg breadth ratio is still around 2:1. The (1996) reviewed the genus, concluding that, ascospores of M. columbiana (G. Merr.) although M. grossa differed from the other Ekman approach those of M. allantoidea in species in several minor respects, they were shape, but they are much smaller (12–)15– best considered as belonging to a single 195–7 m and have a length/breadth ratio variable genus. This view was given some of only c. 3:1 (Noble 1980). support by molecular data (Ekman 2001), Megalaria allantoidea is united with other which showed that M. grossa and M. laureri species of Megalaria by its apothecial (Th. Fr.) Hafellner formed a monophyletic anatomy and pigmentation as described group with high jack-knife support (74– by Ekman & Tønsberg (1996). That is: a 94%), although with rather long branch crustose thallus with biatorine apothecia; length. Ekman’s data also showed that the wide excipulum of radiating hyphae; Megalariaceae should be included in the unbranched paraphyses; hyaline, 1-septate, Ramalinaceae C. Agardh. thick-walled, non-halonate ascospores; More recently, the range of both distri- Biatora-type acus; and green and red pig- bution and substratum of the genus has been ments. The two apothecial pigments present widened by the inclusion of M. jemtlandica in M. allantoidea correspond to those of (Th. Fr. & Almq.) Fryday, which occurs on Ekman & Tønsberg (1996); Cinereorufa- terricolous bryophytes in the arctic (Fryday green corresponding to the first pigment in 2004b), the saxicolous species M. imshaugii their Table 2 (p. 38) and Atra-red to the Fryday and M. macrospora Fryday from second. However, the excipular hyphae Campbell Island (Fryday 2004a), M. beech- of M. allantoidea have inflated cells up to ingii Lendemer from south-eastern U.S.A. 12–15 m across, rather than the much (Lendemer 2007), and several corticolous narrower hyphae typical of the genus. This species from New Zealand (Galloway 2004). character also occurs in M. imshaugii, M. In addition, there are also several other macrospora and a number of other species corticolous and saxicolous species from from the Southern Hemisphere that are South America and the West Indies in either still included in Catillaria (e.g. Catil- MSC that are currently either included in laria obludens (Nyl.) Zahlbr.) or are appar- Catillaria or are apparently undescribed. ently undescribed, and appears to be an All of these species have Bacidia/Biatora- extension of the exciple structure of some type asci (not Lecanora-type as stated by Northern Hemisphere, corticolous species Fryday 2004b for M. imshaugii and M. mac- of Megalaria (e.g. M. laureri), which have rospora) leaving M. grossa in an isolated excipular hyphae with cells up to 5 m position in the genus of which it is the type across. species. However, the decision as to whether to remove the other species to a separate Additional specimens examined. Kerguelen: Presqu’ile Ronarc’h, cliffs on NE face of le Pouce, [c. 49(31.75#S genus, leaving Megalaria as a monospecific 70(10.0#E], 700 m, 1971, G. C. Bratt 71/615 (MSC); genus must await a full molecular analysis of ibid., summit of Mt des Feés, [c. 49(33.0#S representative material. 70(17.0#E], 381m, 1971, G. C. Bratt 71/715, 71/729 The large allantoid ascospores of M. (MSC); Peninsule Courbet, Val Studer, Grande Cascade, c. 800 m SE of Lac Supe´rieur, 49(17#5$S, allantoidea are unique in Megalaria in both 70(03#3$E, 100 m, shaded, humid ravine, overgrowing size and shape, being 40–50 m long and mosses on sloping, exposed basaltic rock, 1998, R. S. having a length/breadth ratio of 5–6:1. Poulsen 388 (C).

I thank the US National Science Foundation (NSF) for Lecanoraceae und Lecideaceae. Beiheft zur Nova Awards No. DBI–9808735 and DBI–0237401 (Alan Hedwigia 79: 241–371. Prather, PI) to Michigan State University that facili- Imshaug, H. A. (1971) Kerguelen Islands botanical tated access to Dr. Imshaug’s extensive collections, and expedition. Antarctic Journal of the United States 6: ﬁnanced the visit of Dr U. Søchting and Mr R. Poulsen 108–110. (Copenhagen) to MSC. I further thank Mr Poulsen and Johnson, A. J., Fryday, A. M. & Prather, L. A. (2005) C for placing his collections at my disposal, and Dr The Michigan State University Herbarium Lichen Brian Coppins (Edinburgh) for correcting the Latin Database. Michigan State University Herbarium: diagnosis. http://www.herbarium.msu.edu/Database. Jørgensen, P. M. (2001) Studies in the lichen family R Pannariaceae X. The lichen genus Protopannaria in 22: Ekman, S. (2001) Molecular phylogeny of the the subantarctic islands. Cryptogamie, Mycologie Bacidiaceae (Lecanorales, lichenized Ascomycota). 67–72. Mycological Research 105: 783–797. Lendemer, J. C. (2007) Megalaria beechingii (lichenized Ekman, S. & Tønsberg, T (1996) A new species of ascomycota), a new species from eastern North Megalaria from the North American west coast, America. Opuscula Philolichenum 4: 39–44. and notes on the generic circumscription. Bryologist Kantvilas, G., Hafellner, J. & Elix, J. A. (1999) 99: 34–40. Tasmidella, a new lichen genus from Tasmania, Fryday, A. M. (2004a) New species and records of with a revised circumscription of the family lichenized fungi from Campbell Island and the Megalariaceae. Lichenologist 31: 213–225. Auckland Islands, New Zealand. Bibliotheca Noble, W. (1980) Catillaria columbiana comb. nov. Lichenologica 88: 127–146. from western North America. Bryologist 83: 71–73. Fryday, A. M. (2004b) A new species of Fuscopannaria Mawson, D. (1934) The Kerguelen Archipelago with a green photobiont, and other taxonomic Geographical Journal 83: 18–27. innovations and new records of lichenized-fungi Meyer, B. & Printzen, C. (2000) Proposal for a stand- from Alaska. Bryologist 107: 173–179. ardized nomenclature and characterization of Fryday, A. M. & Prather, L. A. (2001) The lichen insoluble lichen pigments. Lichenologist 32: collection of Henry Imshaug at Michigan State 571–583. University Herbarium (MSC). Bryologist 104: Poulsen, R. S., Schmitt, I., Søchting, U. & Lumbsch, 464–467. H. T. (2001) Molecular and morphological studies Galloway, D. J. (2004) New lichen taxa and names in on the subantarctic genus Orceolina (Agyriaceae). the New Zealand mycobiota. New Zealand Journal Lichenologist 33: 323–329. of Botany 42: 105–120. Varney, B. M. (1926) Climate and weather at Hafellner, J. (1984) Studien in Richtung einer Kerguelen Island. Monthly Weather Review 54: naturlicheren Gliederung der Sammelfamilien 425–426.

Accepted for publication 20 April 2007