LICHEN GENUS MICAREA IN EUROPE 201 excipular hyphae are conglutinated, broad (c. 2-2-5 /u,m in K), and pachydermatous. It is closely allied to Bacidia subrudecta (Vainio) Zahlbr. and may have affinities with Byssoloma, but I think it is best retained in Bacidia s. lat. pending fur<strong>the</strong>r study. The material distributed by Kalb {Lich. Neotrop. Exs. ^2) as 'Micarea nigrata' is M. lignaria var. lignaria. Psilolechia clavulifera (Nyl.) Coppins, comb. nov. - Lecidea clavulifera Nyl. in Flora, Jena 52: 294 (1869). - Micarea clavulifera (Nyl.) Coppins & P. James, in D. Hawksw. , P. James & Coppins in Lichenologist 12: 107 (1980). Type: Finland, Lapponia kemensis, Muonionska [Muonio], Keimioniemi, ad radicem abietis, 1867, J. P. Norrlin 627 (H - lectotype!; isolectotypes: H!, H-NYL 20855!). Lecidea clavulifera f. subviridicans Nyl. in Flora, Jena 60: 463 (1877). Type: Ireland, West Galway, Connemara, Kylemore, in a cave on <strong>the</strong> NW side <strong>of</strong> Doughraugh mountain, 1876, C. Larbalestier (BMlectotype! Isolectotypes: BM ex K!; also distributed as Larbal. Lich. Herb. 29: BM!, H!). Thallus effuse, whitish or greenish white, <strong>of</strong> dispersed to coalescing, irregular granular-areolae c. Fig. 57 Psilolechia clavulifera (Coppins 3614, E). A, spores. B, paraphyses. C, phycobiont cells (cell contents and mycobiont hyphae omitted). Scale = 10 /xm.
202 BRIAN JOHN COPPINS 0-1-0-2 mm diam. Areolae <strong>of</strong>ten disintegrating to form a scurfy-granular crust. Phycobiont not micareoid; cells in clusters and tightly bound by short-celled hyphae, but haustoria not observed; clusters interconnected by filamentous hyphae c. 1-5-2 ju.m wide. Phycobiont cells irregularly globose, broadly ellipsoid or oblong, c. 5-12(-18)x3-8 />tm, <strong>of</strong>ten arranged in pairs or in short chains <strong>of</strong> up to four cells (Figs. 57C). Apo<strong>the</strong>cia convex-hemispherical and immarginate from <strong>the</strong> beginning, sometimes becoming subglo- bose, more rarely tuberculate, grey-black with bluish tinge, but shade forms sometimes whitish, blue-grey or grey-brown, 0-l-0-3(-0-4) mm diam; base <strong>of</strong> apo<strong>the</strong>cia with a white rim (c. 50 /xm wide) <strong>of</strong> outwardly radiating hyphae. Hymenium 28-35 /xm tall, dilute straw (shade forms), or dilute greenish or aeruginose (K-l- green intensifying) especially in <strong>the</strong> upper part. Asci cylindrical-clavate, 25-30x5-7 /im, 8-spored. Spores ovoid, oblong-ovoid or dacryoid, simple, 3-6x(l-)l-2-l-7(-2) /xm (Fig. 57A). Paraphyses (Fig. 57B) numerous, usually branched, sometimes anastomosing, distinctly septate and appearing ± articu- lated, stout, 1-3-2 /Ltm wide; apices sometimes ± clavate and up to 3 jxm wide, walls not pigmented but <strong>of</strong>ten surrounded by deeply pigmented gel matrix. Hypo<strong>the</strong>cium c. 50-70 /tim tall, hyaline, or dilute greenish or aeruginose (K— , HNO3-I- red) but <strong>the</strong>n never darker than <strong>the</strong> hymenium. Excipulum indistinct, <strong>of</strong> radiating hyphae that protrude as loose, hyaline hyphae c. 1-5-2 /xm wide and up to 50 /x,m long. Pycnidia not found. Chemistry: All parts K- , KC- , C- , PD- ; no substances detected by t.l.c. Preliminary studies <strong>of</strong> Lecidea clavulifera by Mr P. W. James and myself led us to transfer it to Micarea. However, critical studies have given me second thoughts on this placement. The distinct white rim <strong>of</strong> protruding excipular hyphae (superficially like those <strong>of</strong> Byssoloma spp.), stout and distinctly septate paraphyses, and <strong>the</strong> unusual phycobiont are all uncharacteristic <strong>of</strong> a Micarea. The consideration <strong>of</strong> Psilolechia Massal. as an alternative genus for L. clavulifera was prompted by observations <strong>of</strong> a lichenicolous member <strong>of</strong> <strong>the</strong> CaHciales, Microcalicium arenarium (Hampe ex Massal.) Tibell; M. arenarium is usually found as a parasite <strong>of</strong> Psilolechia lucida (Ach.) M. Choisy (see Tibell, 1978), but at three localities in Scotland I have found it on L. clavulifera. At <strong>the</strong> Berwickshire locality <strong>the</strong> host and parasite occurred in abundance on roots, stones, and soil <strong>of</strong> two up-ended trees (Fraxinus); P. lucida was also present in quantity but was not parasitized by M. arenarium. With <strong>the</strong> exception <strong>of</strong> pigmentation, sections <strong>of</strong> <strong>the</strong> ascocarps <strong>of</strong> P. lucida and L. clavulifera show ± identical anatomical features, e.g. nature <strong>of</strong> paraphyses, size and shape <strong>of</strong> asci and spores, and excipulum with numerous protruding hyphae. P. lucida differs from P. clavulifera in <strong>the</strong> yellow-green colour <strong>of</strong> its leprose-granular thallus and apo<strong>the</strong>cia (due to presence <strong>of</strong> pulvinic acid derivatives), and a different phycobiont with ± globose cells 5-14 /xm diam. The thallus hyphae <strong>of</strong> both species are identical in appearance, and pycnidia are not known in ei<strong>the</strong>r species. A recent account <strong>of</strong> P. lucida is given by James (in Poelt & Vezda, 1981). The phycobiont <strong>of</strong> P. clavulifera is very unusual; it is reminiscent <strong>of</strong> <strong>the</strong> Stichococcus phycobiont <strong>of</strong> species such as Chaeno<strong>the</strong>ca stemonea and Coniocybe furfuracea, but its cells are much larger. P. clavulifera occurs in communities <strong>of</strong> <strong>the</strong> Micareetum sylvicolae, on roots, stones, and consolidated soil <strong>of</strong> underhangs on banks or <strong>the</strong> root systems <strong>of</strong> up-ended trees. Most <strong>British</strong> records are from Scotland, but it is also known from Cumbria, south Wales and western Ireland (Map 28). From outside Britain I have seen material <strong>of</strong> it from Finland, Germany, and Czechoslovakia (see below). Additional specimens <strong>of</strong> P. clavulifera examined: <strong>British</strong> Isles. Brecon (V.C.42). 22/94: Upper Dyfnant Valley, 1982, Woods (hb Woods). Cumberland (V.C.70). 35/54: Baron Wood, by River Eden, 60-75 m dry sandstone cliff, 1979, Coppins 4344 (E). Berwick (V.C.81). 36/76: W <strong>of</strong> Elba, S side <strong>of</strong> Whiteadder Water, exposed roots <strong>of</strong> up-ended Fraxinus, 1981, Coppins 8890 (E - Microcalicium arenarium folder). West Perth (V.C.87). 27/40: Aberfoyle, <strong>the</strong> Trossachs, on stone amongst roots <strong>of</strong> up-ended tree, 1978, Coppins3661 (E). Mid Perth (V.C. 88). 27/55: Black Wood <strong>of</strong> Rannoch, stone in bank by track, 1976, Coppins 4654 (E); 27/81: Crieff, Drummond Wood, roots and sandstone <strong>of</strong> up-ended tree, partly parasited by Microcalicium arenarium, 1978, Coppins 3164, 3625 (E). Angus (V.C. 90). 37/33: Sidlaw Hills, Auchterhouse Hill, 396 m, underside <strong>of</strong> boulder, 1975, Coppins 846 (E - Microcalicium arenarium folder). South Ebudes (V.C. 102). 16/49: Colonsay, Coille Mhor, roots and soil <strong>of</strong> up-ended Betula, 1981, Coppins 8878 (E). Mid Ebudes (V.C. 103). Mull: 17/32: Bunessan, Ardfenaig Woods, 1970, James (BM); 17/54: Salen, 1968, James (BM); 17/55; Aros House, roots <strong>of</strong> up-ended tree, 1968, James (BM). West Galway (V.C.H.16). 84/72 (L/65): Connemara, near Clifden, 1878, Larbalestier (BM). Finland. Satakunta: Siikainen, Vuorijarvi, Vaasaneva, sandy soil amongst roots <strong>of</strong> Picea, 1936, Laurila (GZU, H). W. Germany. Baden-Wiirttemberg: Heidelberg, Konigstuhle, on sandstone, 1883, Zwackh (H-NYL p.m. 4229). Czechoslovakia. Slovakia, Nizke Tatry, Liptovska Teplicka, Dzurova, 18 -, Lojka 4289 (BM).
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