Lichenologist 31(3): 231238 (1999) Article No. lich.1998.0202 Available online at http://www.idealibrary.

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A NEW SPECIES OF SCHAERERIA FROM TASMANIA

Gintaras KANTVILAS*

Abstract: The new species, Schaereria bullata Kantvilas from alpine Tasmania, is described and illustrated, with notes on its characterization, distribution and habitat. A key to the seven known species of the genus is provided.
 1999 The British Lichen Society

Introduction In the course of systematic and oristic studies on Tasmanian lichens, an undescribed taxon from the genus Schaereria has been discovered. Schaereria belongs to the monotypic family Schaereriaceae and, in the past, has occupied a rather ambiguous systematic position within the Pezizales (Hafellner 1984; Eriksson & Hawksworth 1985). However, following the detailed study of many similar and related genera by Lunke et al. (1996) and Lumbsch (1997), this family is now included in the Lecanorales, suborder Agyriineae. Other representatives of this suborder present in the Tasmanian lichen ora include Agyrium, Lithographa, Placopsis, Placynthiella, Rimularia, Trapelia and Trapeliopsis, all of which are now included in an expanded concept of the Agyriaceae as dened by Lumbsch (1997)). Problems with the typication of the genus were highlighted by Hawksworth & David (1989). The resolution of these as recommended by the Committee for Fungi and Lichens is given in Gams (1993) and presented below. Materials and Methods
The study is based on the authors collections in HO, and on a range of reference material of non-Tasmanian taxa listed below. Anatomical and morphological data were also derived from published sources. Anatomical observations were undertaken on hand-cut sections mounted in water and a range of standard media: 10% KOH, Lugols iodine and ammoniacal erythrosin. All anatomical measurements are based on sections mounted in water. Routine chemical analyses were undertaken by thin-layer chromatography using standard methods (White & James 1985). Abbreviations of herbaria cited follow Holmgren et al. (1990). Reference specimens examined Schaereria fuscocinerea. Austria: Salzburg: Lungau, Schladminger Tauern, 13201590 m, 8 ix 1981, J. Poelt (GZU, HO).U.S.A.: Colorado: Rocky Mtns, Clear Creek Co., 28452900 m, 4 ix 1977, R. A. Anderson & J. Poelt (H. Hertel: Lecideaceae Exsiccatae Nr 218) (BM).Sweden: Kristineberg: South Skaftn, 15 viii 1967, P. W. James (BM); Torne Lappmark: Kiruna par. Mt *Tasmanian Herbarium, G.P.O. Box 25204, Hobart, Tasmania 7001, Australia. 00242829/99/030231+08 r30.00/0  1999 The British Lichen Society

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Jieprencorru, 750 m, 28 viii 1991, H. Hertel 34592 (H. Hertel: Lecideaceae Exsiccatae Nr 258) (BM). Corsica: Monte dOro-Massiv, Col de Vizzavona, c. 1450 m, 3 xi 1993, H. Mayrhofer & E. Unger (BM).Germany: Bavaria: Oberfranken, road between Pottenstein and Gszweinstein, vi 1863, Arnold, Lich. Exs. 227 (BM). Schaereria cinereorufa: United Kingdom: Glen Fender, Blair Athole, ix 1870, J. M. Crombie (BM); Glen Cove, Glen Fee, 25 vi 1965, P. W. James (BM); Myndd Deulyn, Llyn Crafnant, 75 m, 17 v 1958, P. W. James (BM); Glen Fender, Blair Athole, Meall Daill-mir, 405 m, 27 ix 1970, P. B. Topham (BM); Ben Doran, Argyll, viii 1929, W Watson (BM).Sweden: Uppsala, Th. M. Fries (ex hb. I. Carroll 1874) (BM). Schaereria corticola. Norway: Nord-Trndelag: Grong, 80100 m, 8 viii 1984, T. Tnsberg 8972 (HO).

Taxonomy Schaereria Krber, Syst. Lich. Germ.: 232 (1855). Type: Schaereria lugubris, Falkenstein, Krempelhuber (M) (type cons.) [= Schaereria cinereorufa Schaer.]. The main characters of the genus are a crustose or squamulose thallus, a trebouxioid photobiont, lecideine apothecia with a well-developed, persistent proper exciple and a non-gelatinized hymenium with asci and paraphyses separating easily. The characteristic asci are of the Schaereria-type: cylindrical, with thin, faintly amyloid walls, lacking a tholus, and containing eight simple spores (see illustrations in Hafellner 1984 and Hertel & Zrn 1986). The morphology, anatomy, chemistry and apothecial ontogeny of the genus are discussed in detail by Lunke et al. (1996), and additional details on the genus are provided by Hawksworth (1992), Thomson (1997), Rambold (1989), Clauzade & Roux (1985), Hertel & Zrn (1986), Hafellner (1984) and Tnsberg (1992). Following the work of Lumbsch (1997), seven species are now included in Schaereria: S. cinereorufa Krb., S. corticola Muhr & Tnsberg, S. fabispora Hertel & Zrn, S. fuscocinerea (Nyl.) Clauzade & Cl. Roux, S. parasemella (Nyl.) Lumbsch, S. xerophila H. Mayrhofer & Rambold and the new taxon described below. Most species are found primarily in cool to cold climates of the Northern Hemisphere, but S. fuscocinerea is also bipolar whereas S. xerophila occurs in lowland Queensland. Species of Schaereria are typically saxicolous on siliceous rocks although S. corticola is epiphytic and S. parasemella is lichenicolous. Thus the occurrence of a new taxon in the mountains of Tasmania is consistent with the global distribution pattern of the genus as a whole. It also accords with the predominantly alpine ecology of the other genera of the Agyriineae in Tasmania. Schaereria bullata Kantvilas sp. nov.
Species thallo bullato, granulari vel squamuloso, apotheciis supercialibus, haud immersis, et sporis globosis, Schaereriae cinereorufae Krb. maxime similis sed epithecio brunneo, non aeruginoso, excipulo non opaco et prothallo carenti praecipue diert. Typus: Australia, Tasmania, Legges Tor, 41 32 S, 147 39 E, on skeletal soil or directly on at rock surfaces in alpine boulder eld, 1570 m altitude, 28 April 1998, G. Kantvilas 103/98 (HOholotypus).

(Fig. 1)

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F. 1. Schaereria bullata (holotype). A, habit of thallus and apothecia; B, asci, ascospores and paraphyses, with amyloid parts stippled. Scales: A =05 mm; B =10 m.

Thallus dark brown to grey-brown, occasionally reddish brown, saxicolous, forming scattered irregular patches c. 1020 mm wide over skeletal soil or bryophytes, squamulose and composed of crowded, coalescing granules; squamules convex to bullate, 0108(1) mm wide, 0205 mm thick, lacking a cortex but with an outer brownish layer 518 m thick, sometimes overlain by an epinecral layer up to 8 m thick; photobiont cells irregularly roundish, 813 m wide. Apothecia typically abundant, scattered or clustered, roundish, sometimes rather irregularly so due to mutual pressure, 051 mm wide when mature, 02505 mm tall, supercial, constricted at the base, never immersed or adnate; proper margin prominent, persistent, inrolled, dark brown to black, sometimes concolorous with the thallus, 02025 mm thick; disc persistently plane, concolorous with the margin or a little darker, smooth to verruculose when old. Exciple in section cupular, 55140 m thick at the sides, c. 80 m thick at the base below the hymenium, pale brown to brown within, with a dark brown or reddish brown outer edge, composed mainly of irregularly roundish cells, 69 m wide; pigment becoming rather grubby grey-brown in KOH. Hymenium colourless, 85110 m thick, separating very easily in water, with a pale reddish brown or yellowish brown epithecial layer up to c. 12 m thick, becoming grey-brown in KOH. Hypothecium mainly colourless, at least in the upper part, pale yellow-brown to brown below, 100160 m thick. Asci narrowly cylindrical, 8095 1013 m; walls I+pale blue. Paraphyses cylindrical, simple or occasionally forked towards the apices, 1525 m thick, typically narrower in the lower part and wider above; apices tapered or, more commonly, slightly swollen to 3545 m thick, often moniliform, colourless or faintly brownish pigmented. Ascospores spherical,

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F. 2. Distribution of Schaereria bullata.

strictly uniseriate, non-halonate, (6)811(115) m in diam., typically with a thin but distinct wall c. 05 m thick. Pycnidia immersed, ask-shaped, 80115 m wide, without pigmented walls, visible as minute black dots in the upper surface of the thallus. Conidia narrowly cylindrical, simple, 36 051 m. C Chemistry. Containing two unknown substances; cortex and medulla K , KC , CK , PD , UV . ,

Distribution and ecology in Tasmania. Schaereria bullata is a rather rare species, restricted mainly to Tasmanias dolerite alpine provenance (see Kantvilas 1995). It is known mainly from the north-eastern highlands and Cradle Mountain, where it may be locally abundant, with a single outlying occurrence on Ordovician conglomerate on the West Coast Range (Fig. 2).

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Lichens with a similar Tasmanian distribution include Cetraria australiensis, Cladonia bimberiensis, C. ecmocyna, Protoparmelia badia and Umbilicaria subglabra. It has been recorded above 900 m altitude in boulder elds, scree slopes and clis. The new species has a distinctive ecology, occurring on horizontal rock plates where moisture accumulates temporarily and a thin skin of soil develops. It thus grows in an intermediate niche between the extremely exposed rock surfaces where crustose lichens, Stereocaulon caespitosum, Pseudephebe pubescens and other typical and common alpine lichens are found, and the moister sheltered rock crevices colonized by bryophytes, Siphula complanata, Psoroma hypnorum, basal squamules of Cladonia, and other incipient macrolichens. Schaereria bullata is rather easily recognized in the eld, but supercially, it can resemble some smaller members of the Pannariaceae that may occur in similar habitats, for example, Parmeliella ligulata, Degelia neozelandica and Psoroma hypnorum. This is especially so because its apothecia have a thick, dark margin that is often concolorous with the thallus and may appear to be lecanorine. The granular thallus may also resemble that of Catillaria contristans or some Micarea species, taxa also sometimes associated with S. bullata, but these have convex apothecia lacking a distinct margin. Remarks. The overall habit, spore-type and saxicolous montane ecology of S. bullata ally this species most closely to the widespread Northern Hemisphere taxon, S. cinereorufa, but the two dier by several consistent, albeit subtle characters. The most striking dierence is the complete absence in the new species of any blue-green pigmentation (intensifying greenish in KOH) that occurs in the epithecium, upper part of the hymenium, upper edge of the exciple and around the ostiole of the pycnidia of S. cinereorufa. Schaereria cinereorufa diers further by its generally darker apothecia, manifested anatomically by an opaque, black-brown exciple in section. Although some authors describe the hypothecium of S. cinereorufa as being brown to blackish, the pigmented tissue is actually a cupular exciple that extends beneath the hymenium and a colourless or pale hypothecium. Such an exciple (as distinct from a heavily gelatinized cupular hypothecium) is found in several unrelated groups and the taxonomic signicance of this feature is unclear (Feige & Lumbsch 1998). In gross morphology, S. bullata falls within the range of variation displayed by S. cinereorufa, although it never develops a microphylline habit sometimes seen in well-developed forms of the latter, nor does it have a black prothallus. The chemistry of both S. cinereorufa and S. bullata is unresolved and includes unknown compounds, one of which is common to both species. However, a qualitative dierence between the species has been observed, with the former also containing gyrophoric acid (sometimes only in trace amounts), whereas the latter has a second unidentied compound (Fig. 3). Another species closely related to S. bullata is the Queensland taxon, S. xerophila. Unfortunately, the type material could not be located for the present study, but the description and illustration in Rambold (1989) is comprehensive. Both taxa have a brown epithecium, internally unpigmented exciple

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F. 3. Standard TLC analyses in solvent systems TDA and G (after White & James 1985) of diagnostic compounds in Schaereria. 1: reference; 2: S. bullata (Kantvilas 103/98); 3: S. cinereorufa (Watson s.n.); 4: S. fuscocinerea (Lecid. Exsicc. 258). Compounds: A: atranorin; B: norstictic acid; C: unknown 1; D: gyrophoric acid; E: unknown 2.

and globose, non-halonate spores. However, S. xerophila has an areolatecrustose thallus, subimmersed apothecia and a completely disjunct ecology, occurring on lowland, inland rocks at latitude 25 S. The dierences between S. bullata and the remaining species in the genus are summarized in the identication key (below). These are based on combinations of thallus and spore morphology, apothecial pigmentation and thallus chemistry. The only other species recorded in the Australasian region (Rambold 1989; Hertel 1989) is S. fuscocinerea, which contains gyrophoric acid and has a crustose-areolate thallus, adnate to immersed apothecia, mostly <05 mm diameter, a blue-green epithecium and ellipsoid, biseriate spores.
Specimens examined: Australia: Tasmania: Mt Saddleback summit area, 41 24 S, 147 45 E, 1200 m, 1993, G. Kantvilas 92/93 (HO); Little Horn, 41 41 S, 145 58 E, 1320 m, 1995,

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G. Kantvilas 88/95 (HO); Mt Victoria, 41 20 S, 147 50 E, 1000 m, 1997, G. Kantvilas 7/97 (HO); western anks of Legges Tor, 41 32 S, 147 39 E, 1530 m, 1998, G. Kantvilas 108/98 (HO); western slopes of Mt Geikie, 41 58 S, 145 34 E, 900 m, 1998, G. Kantvilas 198/98 (HO).

Key to the species of Schaereria 1. 2(1) Ascospores globose . . . . . . . . . . . . . . . . . . . . . . . . . 2 Ascospores ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . 4 Epithecium blue-green; exciple opaque black-brown; found in montane and cold habitats of the Northern Hemisphere . . . . . S. cinereorufa Epithecium brown; exciple internally pale brown to brown, not opaque; occurring in the Southern Hemisphere . . . . . . . . 3 Thallus crustose-areolate; apothecia semi-immersed; in lowland Queensland . . . . . . . . . . . . . . . . . . . . . S. xerophila Thallus granular to squamulose; apothecia supercial, not immersed; alpine Tasmania . . . . . . . . . . . . . . . . . . . . S. bullata Thallus very thin to absent; spores halonate . . . . . . . . . . . . 5 Thallus crustose-areolate; spores lacking a halo . . . . . . . . . . 6 Thallus corticolous, sorediate; occurring in the temperate Northern Hemisphere . . . . . . . . . . . . . . . . . . . . . . S. corticola Thallus lichenicolous on Biatora; known only from Finland . . . . S. parasemella Spores ellipsoid, straight, mostly ^16 m long; bipolar, widespread in cold, montane environments . . . . . . . . . S. fuscocinerea Spores reniform, >16 m long; known only from Norway. . . . . . S. fabispora Lichenicolous fungus In general, the genus Schaereria can be a common host for lichenicolous fungi and lichens. Scheidegger (1987) records two species of Buellia, Triebel (1989) records species of Endococcus and Muellerella, whilst Rambold & Triebel (1992) record species of Buellia, Miriquidica and Rhizocarpon, all from Schaereria tenebrosa (= S. fuscocinerea). One Tasmanian specimen of S. bullata is infested with a species of Dactylospora belonging to the D. australis-D. saxatilis group. This taxon is characterized by the following: apothecia up to 025 mm diam., c. 140 m thick, excipulum brown, opaque, c. 1525 m thick, hymenium 5085 m thick, epithecium brown, 818 m thick, hypothecium 6070 m thick, reddish brown, spores brown, 1-septate, 913 57 m, paraphyses with apices capitate, pigmented, 3545 m thick. According to Dr D. Triebel (in litt.), this taxon may well be new, diering from D. australis by the somewhat broader and longer spores, and from D. saxatilis by the slightly darker hypothecium and thinner, indistinctly radiate exciple. Further collections are required to establish whether these dierences are consistent.

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Specimen examined: Australia: Tasmania: western anks of Legges Tor, 41 32 S, 147 39 E, 1530 m, 1998, G. Kantvilas 108/98 pp (HO, M). This project was undertaken at the Natural History Museum, London; I am grateful to the Keeper and sta for providing the facilities for carrying out the work, and to members of the Lichen Section, especially Mats Wedin, Pat Wolseley and William Purvis, for their assistance and support during the course of my visit. I also thank Dagmar Triebel for her speedy comments on the lichenicolous parasite, and Jean Jarman for assistance with the gures. R Clauzade, G. & Roux, C. (1985) Likenoj de Okcidenta Europo. Illustrita Determinilibro. Bulletin de la Socit Botanique du Centre-Ouest, Nouvelle Srie, Numro Spcial 7. Eriksson, O. & Hawksworth, D. L. (1985) Outline of the Ascomycetes-1985. Systema Ascomycetum 4: 177. Feige, B. & Lumbsch, H. T. (1998) The ascomata development in Arctopeltis thuleana (Lecanoraceae) and its systematic signicance. Cryptogamie, Bryologie-Lichnologie 19: 147153. Gams, W. (1993) Report of the committee for fungi and lichens: 3. Taxon 42: 112118. Hafellner, J. (1984) Studien in Richtung einer natrlicheren Gliederung der Sammelfamilien Lecanoraceae und Lecideaceae. Beihefte zur Nova Hedwigia 79: 241371. Hawksworth, D. L. (1992) Schaereria Th. Fr. (1885). In The Lichen Flora of Great Britain and Ireland (O. W. Purvis, B. J. Coppins, D. L. Hawksworth, P. W. James & D. M. Moore, eds). 557558. London: Natural History Museum Publications. Hawksworth, D. L. & David, J. C. (1989) 933944 Proposals for nomina conservanda and rejicienda for ascomycete names (lichenized and non-lichenized). III. Taxon 38: 493499. Hertel, H. (1989) New records of lecideoid lichens from the Southern Hemisphere. Mitteilungen der Botanischen Staatssammlung Mnchen 28: 211238. Hertel, H. & Zrn, L. (1986) Schaereria fabispora (Ascomycetes lichenisati)- eine neue Art aus Norwegen. Mitteilungen der Botanischen Staatssammlung Mnchen 22: 477483. Holmgren, P. K., Holmgren, N. H. & Barnett, L. C. (1990) Index Herbariorum Part I: The Herbaria of the World. 8th Edn. New York: New York Botanic Gardens. Kantvilas, G. (1995) Alpine lichens of Tasmanias South-West wilderness. Lichenologist 27: 433449. Lumbsch, H. T. (1997) Systematic studies in the suborder Agyriineae (Lecanorales). Journal of the Hattori Botanical Laboratory 83: 173. Lunke, T., Lumbsch, H. T. & Feige, G. B. (1996) Anatomical and ontogenetic studies on the lichen family Schaereriaceae (Agyriineae, Lecanorales). Bryologist 99: 5563. Rambold, G. (1989) A monograph of the saxicolous lecideoid lichens of Australia (excluding Tasmania). Bibliotheca Lichenologica 34: 1345. Rambold, G. & Triebel, D. (1992) The inter-lecanoralean associations. Bibliotheca Lichenologica 48: 1201. Scheidegger, C. (1987) Buellia uberior und B. miriquidica (Physciaceae, Lecanorales), zwei lichenicole Krustenechten auf Schaereria tenebrosa. Botanica Helvetica 97: 99116. Thomson, J. W. (1997) American Arctic Lichens. 2. The Microlichens. Madison: University of Wisconsin Press. Tnsberg, T. (1992) The sorediate and isidiate, corticolous crustose lichens in Norway. Sommerfeltia 14: 1331. Triebel, D. (1989) Lecideicole Ascomyceten. Eine Revision der obligat lichenicolen Ascomyceten auf lecideoiden Flechten. Bibliotheca Lichenologica 35: 1278. White, F. J. & James, P. W. (1985) A new guide to microchemical techniques for the identication of lichen substances. Bulletin of the British Lichen Society 57 (suppl): 141. Accepted for publication 5 December 1998