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LICHEN GENUS MICAREA IN EUROPE 143 Paratype: Germany: 'Ibbenbiiren in Westfalen, Lahm, Hb. Hepp.', on sandstone (BM!). Stereocauliscum gomphillaceum Nyl. in Flora, Jena, 48: 211 (1865). - Micarea gomphillacea (Nyl.) Vezda in Folia geobot. phytotax., Praha 5: 321 (1970). - Bilimbia milliaria f. [as 'f?'] gomphillacea (Nyl.) Blomb. & Forss., Enum. PI. Scand.: 82 (1880). - Micarea lignaria [as 'ligniaria'] f. gomphillacea (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 94 (1892). Type: Finland, Tavastia australis, HoUola, Tiirismaa, Pirunpesa, [on granite rock], 1863,/. P. Norrlin (H-NYL 40217 -lectotype! [t.l.c: argopsin];isolectotypes: H-NYL 40214!, 402151,40216!, 40218! and p.m. 631!). Lecidea sabuletorum var. milliaria f. nigrata Nyl. in Not. Sdllsk. Fauna Fl. fenn. Forh. 8: 151 (1866). Type: USSR, Kola Peninsular, Lapponiaponojensis,Ponoi, 1863, M /. Fe/Zman (H-holotype!;H-isotype!). See note (i) below. Lecidea milliaria var. triseptata Nyl. in Lamy in Bull. Soc. bot. Fr. 25: 441 (1878). - Bilimbia triseptata (Nyl.) Arnold in Flora, Jena 67: 572 (1884), non Bacidia triseptata (Hepp in ZoU.) Zahlbr. Type: France: Haute Vienne, 'Sur des roches entre Chalard et St Junien, Lamy (H-NYL 18367! - lectotype, as 'L. milliaria var. 3-septata Nyl.'). See note (ii) below. Lecidea meizospora Harm, in Bull. Seanc. Soc. Sci. Nancy II, 33 (1898): 63 (1899). - Bacidia meizospora (Harm.) Zahlbr., Cat. lich. univ. 4: 122 (1926). Type: France, Vosges, Docelles, on Calluna roots, V. and H. Claudel & F. J. Harmand (ANGUC - lectotype!). Lecidea trisepta var. polytropoides Vainio in Ark. Bot. 8 (4): 106 (1909). Type: USSR, Magadanskya Oblast (N.E. Siberia), 'Pitlekai, peninsula Jinretlen' [c. 67°7'N 173°24'W], on plant debris, 1878-9, E. Almquist (TUR-VAINIO 21274, fertile part- lectotype!). See note (iii) below. Patellaria milliaria var. xylophila Wallr., Fl. crypt. Germ. 1: 349 (1831); nom. inval. (Art. 26). Lecidea milliaria var. saxigena Leighton, Lich. Brit. Exs. 210 (1856); nom. nudum (Art. 32); see note under M. peliocarpa. Type: England, Shropshire, Neescliff [Nesscliff] Hill, Leighton, Lich. Brit. Exs. 210 (E- lectotype!; isolectotypes: BM! DBN! MANCH! NMW!). Notes: (i) The type material mainly consists of the black stroma of Bryomyces gymnomitrii Dobb. on Gymnomitrium sp. , with only a few apothecia of M. lignaria present. (ii) Lamy (loc. cit.) gives the locality as 'entre le Chalard et Saint-Yrieix'. The label on the lectotype specimen is entirely in Nylander's handwriting and there may have been an error in transcription. (iii) Vainio's diagnosis included the description of soralia. A small portion of the type specimen is a sterile sorediate thallus (C- , PD- phycobiont cells 7-14 /am diam) but it is not a Micarea and is not part of ; the larger, fertile portion. The latter belongs to Micarea lignaria and is chosen as lectotype. Thallus effuse, partially endoxylic or endocuticular, but more usually developed superficially as convex to ± globose areolae. Areolae c. 80-250(-300) /xm diam, discrete, scattered to contiguous, sometimes coalescing to form an uneven crust, grey-white and ± glossy to grey-green or bluish grey and matt, sometimes brown-grey due to the ramification through their surface of brown, toruloid hyphae (cells c. 5-7x4-5-7 /xm) belonging to an unknown fungus. Areolae in section, with a hyaline amorphous covering layer (c. 4—12 /xm tall); walls of hyphae in outermost (exposed) parts often blue-green (K-, HNO3+ red); algal layer usually in direct contact with substratum, but in larger areolae the algal layer may be c. 90-135 ixva tall lying above a medulla. Medulla, when present, up to c. 100 ^im tall, consisting of loosely interwoven hyphae c. 1-1-5 /xm wide, intermixed with dead algal cells and debris derived from the substratum. Phycobiont mic2iVQO\d, cells c. A-1 /xm diam. Apothecia scattered, more usually numerous and crowded, often confluent, immarginate, convex-hemispherical, soon becoming constricted at the base and hence ± globose, occasionally becoming stipitate with a grey-black 'stipe' up to 1 mm tall ('f. gomphillacea''), black, matt or slightly glossy, rarely blue-grey (shade forms), 0-15-0-6(-0-9) mm diam. Hymenium 50-75 /xm, often not sharply delimited from the hypothecium, dilute olivaceous or aeruginose-green (K- HNO3-I- red) in upper, and sometimes lower, part(s) (the middle part often being ± hyaline); in the uppermost part (epithecium) the pigment is more dense and is present not only in the gel matrix but also in the walls of the paraphyses; in reflexed parts of the hymenium towards the base of the apothecium the green pigment is often replaced by a dilute brown pigment; minute (less than 2x1 yu,m) granules of violet (K+ intense aeruginose) pigment are sometimes present and lightly scattered through the hymenium. Asci clavate, 45-50x11-19 /xm. Spores fusiform, straight or slightly curved, (0-)3-7 septate, 16-36(-38)x4-6(-7) /xm. Paraphyses numerous, simple or slightly branched above, but more richly branched in the transition zone between the hymenium and excipulum, sometimes anastomosing, 1-3-1-8 /xm wide; apices often slightly ,
144 BRIAN JOHN COPPINS incrassate and coloured with green or greenish brown pigment and then up to 3 /xm wide, usually coherent (even in K) and, together with the surrounding pigmented matrix, form an epithecium. Hypothecium c. 100-230(-350) /xm tall, dilutely pigmented with pigment confined to gel matrix; upper part dull aeruginose or olive-brownish K- or + dulling, HNO3+ reddish); lower part often without greenish tinge, and then being dilute brownish or ± hyaline; hyphae interwoven or some ± vertically orientated near the hymenium, c. 1-1-7 /am wide; ascogenous hyphae with short, swollen cells c. 2-5-4 /xm wide. Excipulum ± distinct in sections of young, hemispherical apothecia, but soon becoming strongly reflexed and not sharply delimited from the hymenium, dilute brown or sometimes darkish brown along the outer edge. Hypothecial and excipular tissues sometimes elongated vertically to form a stipe ('f. gomphillaced') Pycnidia inconspicuous, ± immersed, with walls green (K-, HNO3-I- red) in exposed upper parts and ± hyaline in immersed lower parts; of three types [only type (c) is common]: (a) c. 100 ixm diam; conidia {macroconidia) curved or hamate, 0-3-septate, 16-22 Xc. 1 /xm; (b) c. 100-140 /xm diam; conidia (mesoconidia) ± cylindrical, oblong-ellipsoid, obovoid or oblong-obovoid, usually distinctly truncate at proximal end, often 2-3-guttulate, 4-7(-7-6)xl-2-l-8 fxm; (c) c. 40-50 ^im diam; conidia (microconidia) narrowly cylindrical, only faintly truncated at proximal end, eguttulate, (4-5-)5-7(-8)x 0-8-1 /xm. Chemistry: Thallus K-, C-, KC-, PD-I- red; sections of apothecia C-, PD-; t.l.c: argopsin. Observations: Micarea lignaria is characterized by its whitish to grey, convex to ± globose areolae which are PD+ red (argopsin; but see var. endoleuca), black, markedly convex to ± globose apothecia, green upper hymenium, dilute greenish or dilute olive-brownish hypothecium, '^nd 3-7-septate, fusiform spores. When on lignum the thallus is often reduced to small scattered areolae and sometimes it is ± entirely endoxylic. The apothecia are less variable in appearance, and pale (shade) forms are very rare. However, the apothecia are occasionally found (especially on rock in dry underhangs) to be stipitate with a 'stipe' (composed of vertically extended hypothecial and excipular tissues) up to 0-4 mm, or even 1 mm, tall (Fig. 3E). These forms have been ascribed the status of form, variety, species and even genus, viz.: Nylander's new species and genus Stereocauliscum gomphillaceum. In the type material of this, some 'stipes' are extremely tall (up to 1 mm) and composed of vertically proliferating apothecia, with the apical apothecia being the youngest; the apothecia are immature (or arrested in their development) with few asci and spores. In my opinion these stalked forms of A/, lignaria result from abnormal development in response to adverse environmental conditions and are, there- fore, not worthy of taxonomic recognition at any rank. M. lignaria is often confused with M. cinerea and M. peliocarpa, but these two species have usually more flattened and often paler apothecia, more richly branched paraphyses, and a hyaline hypothecium. In addition, they contain gyrophoric acid, resulting in the C-l- orange-red (quickly fading) , PD — reactions of their thalli and apothecia. M. ternaria {q. v. ) is very similar to M. lignaria, but has more flattened apothecia with a more discernible excipulum and spores which are never more than 3-septate; furthermore, it lacks any lichen substances. Habitat and distribution: M. lignaria occurs on a wide range of substrata, but is most common in upland districts, growing over bryophytes or plant debris on siliceous rocks and walls, or on exposed peaty ground. Associated lichens found in such habitats in Britain include Arthrorhaphis citrinella, Baeomyces rufus, Cladonia coccifera, C. floerkeana, C. squamosa, C. subcervicornis, Coelocaulon aculeatum s. lat., Lecidea granulosa, L. icmalea, Lecidoma demissum, Lepraria neglecta, Micarea leprosula, M. peliocarpa, Ochrolechia frigida, Parmelia saxatilis, and Pseudephebe pubescens. It often grows over bryophyte mats that are heavily invaded by gelatinous algae, and in such situations in Scotland it has been found with the rare, or overlooked, Arctomia delicatula and Belonia incarnata. When growing directly on rock M. lignaria is mostly confined to sheltered, shaded situations and is sometimes present in the Micareetum sylvicolae in rock underhangs. It is usually present in abundance in the old lead and zinc mine workings of the Pennines and Scotland, where it grows over decaying bryophytes and .
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24 BRIAN JOHN COPPINS (ii) Goniocys
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34 BRIAN JOHN COPPINS Fig. 8 M. ala
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36 BRIAN JOHN COPPINS Fig. 10 A, M.
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38 BRIAN JOHN COPPINS Fig. 12 A,M.
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40 BRIAN JOHN COPPINS r\ U Fig. 14
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44 BRIAN JOHN COPPINS Fig. 18 M. le
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46 BRIAN JOHN COPPINS Fig. 20 A-B,
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48 BRIAN JOHN COPPINS Fig. 22 M. me
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52 BRIAN JOHN COPPINS Fig. 26 M. pr
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54 BRIAN JOHN COPPINS VJ Fig. 28 A,
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56 BRIAN JOHN COPPINS Fig. 30 M. su
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58 BRIAN JOHN COPPINS Fig. 31 - con
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60 BRIAN JOHN COPPINS Fig. 33 M. tu
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64 BRIAN JOHN COPPINS Table 1 Conid
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68 BRIAN JOHN COPPINS Fig. 37 M. ad
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70 BRIAN JOHN COPPINS s o o c o c o
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72 BRIAN JOHN COPPINS n 1/ [\ f] n
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76 BRIAN JOHN COPPINS Fig. 45 M. li
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78 BRIAN JOHN COPPINS B OOQog Fig.
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90 BRIAN JOHN COPPINS fM. elachista
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conferanda (Lecidea) 205 confusula
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populina (Lecidea, Micarea) 31,32 p
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