Bulletin of the British Museum (Natural History)

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LICHEN GENUS MICAREA IN EUROPE 151 Biatora stizenbergeri Hepp, Flecht. Eur. no. 504 (1860). Type: Switzerland, Rifferschwell, on dry plant debris amongst roots in peat moor, Hegetschweiler (BM - lectotype!; isotypes distributed in Hepp. Flecht. Eur. 504: BM!, E!, WRSL!). Lecidea ilyophora Stirton in Scott. Nat. 5: 220 (1879). Type: Scotland, Perthshire, Black Wood of substances detected]). Rannoch, on lignum, ix 1879, J. Stirton BM - lectotype! [t.l.c. : no Lecidea melaena f. catillarioides Vainio in Medd. Soc. Fauna Fl. fenn. 10: 12 (1883). - Micarea melaena f. catillarioides (Vainio) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 83, 96 (1892). Type: Finland, Karelia borealis, Nurmes, Riihivaara, on bryophytes on rock, 1875, Vainio (TUR-VAINIO 21478 -holotype!). Lecidea melaena f. endocyanea Vainio in Medd. Soc. Fauna Fl. fenn. 10: 12 (1883). - Micarea melaena f. endocyanea (Vainio) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill 18 (3): 83, 96 (1892). Type: USSR, Karelia keretina, Kivakka, on mosses on rocks, 1877, Vainio (TUR-VAINIO 21476 - hole- type!). Bilimbia melaena f. aeruginosa Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 254 (1922). Type: Finland, Tavastia australis, Hollola, on lignum, 1875, Vainio (TUR-VAINIO 21461 - lectotype!). Bilimbia melaena f. epiphaeotera Vainio in Acta Soc. Fauna Fl. Fenn. 53 (1): 255 (1922). Type: Finland, Karelia borealis, Nurmes, Louhivaara, 'Kallion syrjalla', on bryophytes and old Cladonia squamules, 1875, Vainio (TUR-VAINIO 21472- lectotype!). ?Bilimbia melaena var. alnicola Savicz in Izv. imp. S. Peterb. bot. Sada 14 (I, Suppl.): 53 (1914). Type: USSR (?LE); request for loan received no reply. Bilimbia milliaria y. [var.] saprophila Korber, Parerg. lich. : 171 (1860); nom. superft. (Art. 63). Thallus effuse, superficial, of small, scattered to coherent or clustered, granular areolae (20-)25-80(-100) /xm diam, sometimes forming an uneven crust up to c. 150 fxm thick. Areolae very variable in colour, pale buff or pale dull green to dark grey-green; in section without an amorphous covering layer, outermost hyphae often with dark green (K-, HNO3-I- red) walls. Thallus often scurfy granular and blackish (especially in damp situations) owing to its disruption by invading dematiaceous fungi and foreign algae; in dry situations (e.g. undersides of fallen trunks) it may have a whitish or pale yellowish , farinose appearance due to a thick covering of an epiphytic alga, each cell of which is coated by hyaline, crystalline incrustations. Phycobiont micareoid, cells c. 4-7 jxm diam. Apothecia numerous, immarginate, convex-hemispherical to ± globose, sometimes tuberculate, black, matt or slightly glossy, 0- 12-0-4 mm diam, or to 0-5 mm diam when tuberculate. Hymenium 30-40(-50) /xm tall, dull to bright aeruginose or grey-blue, K— or K-l- green intensifying; alternatively lower part (or rarely the entire hymenium) dilute purple-brown, K-l- greenish or K-l- purple intensifying; pigment(s) often more dense around the apices of the paraphyses, thus appearing as a dark 'epithecium' (c. 2-10 jxm tall) that contrasts with the more dilute colouration of the rest of the hymenium. Asci clavate, c. 30-40-12-15 jxm. Spores ovoid-oblong or oblong, with rounded apices, straight, (l-)3(-5)-septate, 12-21 x4-5(-5-5) /xm. Paraphyses numerous, branched and anastomosing, sometimes a few unbranched, thin, 0-8-1 ^im wide; apices not swollen, or sometimes slightly incrassate and up to 1-5 jxm wide, hyaline but often individually surrounded by a pigmented hood of dense gel (easily detached in mounts in K, by tapping the cover slip). Occasionally intermixed with ordinary paraphyses are a few which are simple , stout (c. 1 -5-2 jxm wide) and coated with pigment throughout their length so as to appear 3-4 wide. Hypothecium c. 80-160 ^im tall, dark and ± black in thick sections, pigmentation variable (see 'observations' below), usually purple-brown and K— or K-l- purple intensifying, or, outer (more rarely whole) part K-l- olivaceous with central part ('core') remaining ± purplish brown; all parts HNO3-I- purple-red; hyphae c. 0-7-1-5 jxm wide but their walls coated with dense pigment so that they appear to be c. 3-4 /xm wide, interwoven or becoming vertically orientated towards the hymenium, sometimes a few extending into the hymenium as stout pigmented paraphyses; ascogenous hyphae similarly pigment, short-celled, c. 2-5 /am wide. Excipulum indistinct, sometimes evident in sections of young apothecia, green or purplish (usually darker than hymenium, but paler than hypothecium); hyphae radiating, branched and anastomosing, c. 0-7-1 jxm wide. Pycnidia rare, black, of two types: (a) c. 100-140 /xm diam, sessile; walls purple-brown or dull olive, K-l- green; conidia {macroconidia) curved or hamate, 0-7-septate, 18-33 x 1-1-5 /xm; (b)
152 BRIAN JOHN COPPINS c. 40-60 /Ltm diam, semi-immersed to sessile, walls green, K—;conidia (microconidia) fusiform- cylindrical, straight, 4-8-7x0-8-l /itm. Chemistry: Thallus K— , C+ red or C— , PD; sections of apothecia C— ; t.l.c: gyrophoric acid (healthy thalli), or no substances or gyrophoric acid in trace amounts (poorly developed or disrupted thalH). Observations: Micarea melaena is one of the commonest species of the genus, especially in northern Europe, and it displays a considerable amount of variation in thallus appearance and internal apothecial pigmentation. However, my examinations of over 200 specimens have revealed many intermediates between the known extremes. When well-developed, the thallus is composed of cohering, small, granular areolae and is pale buff to dull green in colour. In exposed situations the thallus often becomes a dark green-black due to the more intense pigmentation of the outermost hyphae of the granules. The appearance of the thallus can also be affected by the presence of other organisms (see description and p. 00). When on lignum, in dry exposed situations the thallus may become much reduced to a fine scattering of tiny (less than 30 /xm diam), blackish granules amongst the wood fibres. Such forms proved to be very common, during my explorations in Sweden. M. melaena also exhibits much variation with regard to the colouration of its apothecial tissues (especially the hypothecium). This is due to the presence of three (probably biosynthetically related) pigments, varying in their relative amounts and distribution. These pigments, all of which react HNO3+ purple-red, can be roughly identified as pigment A: green, K-l- green intensifying; pigment B: purple, K+ green; pigment C: purple, K+ purple intensifying. In the epithecium and hymenium pigment A is often found alone, but sometimes it is replaced in the lower hymenium (or in extreme cases throughout the entire hymenium and epithecium) by pigment C. Pigment C is usually dominant in the hypothecium, but is often replaced in the upper and outer parts (or more rarely ± throughout) by pigment B. If thin sections of apothecia in which pigment B is dominant are mounted in K the lower central ('core') of the hypothecium retains a purplish tinge (the rest having turned green), indicating the presence of a small amount of pigment C. The same type of variation, presumably involving identical (or very similar) pigments, is found in several other species, e.g. M. crassipes and M. sylvicola. The pycnidia of M. melaena are very inconspicuous and apparently sparingly produced. However, a close scrutiny of about 100 specimens revealed ten collections with the microconi- dial state and only two (e.g. Coppins 6041) with the macroconidial state. A mesoconidial state was not found but may well occur. Habitat and distribution: M. melaena is widely distributed and especially common in northwest Europe. Although, for a Micarea, it is a conspicuous species, it is often overlooked as a black or charred stain on the tops of stumps and peaty turf! It is a common inhabitant of stumps and fallen decorticate trunks in upland woodlands, or in lowland woodlands on very acid, nutrient-poor soils, and can exist in shaded to extremely exposed, sun-baked situations. In upland woods it can be found on the corticate trunks of healthy trees (especially Betula, Quercus, and Pinus). In areas heavily polluted by SO2 and its derivatives (e.g. West Yorkshire conurbation) it has been found on the acidified bark of Acer and Ulmus. Occurrences on decaying timber work (e.g. fence posts, gate rails and shingles) are not uncommon, especially in upland districts. Lichens associated with M. melaena on lignum and bark in the British Isles include Bryoria capillaris, B. fuscescens, Cladonia spp., Hypocenomyce scalaris, Hypogymnia physodes, Lecidea granulosa agg., L. icmalea, Micarea denigrata, M. prasina, Mycoblastus sterilis, Ochrolechia turneri, Parmelia saxatilis, Parmeliopsis spp., Platismatia glauca, Trapelia corticola, and Usnea spp. M. melaena is often terricolous, growing on oligotrophic, peaty soils, in montane regions or lowland heaths. In the English lowlands it is recorded from heaths in the Sussex Weald and east Norfolk. It is common on ± bare peat in the Pennines and Scottish highlands, but in the latter it is usually replaced near the higher summits (over c. 1000 m) by M. turfosa. Associated lichens on peaty turf in Britain include 'Botrydina' (i.e. Omphalina spp.), Cladonia coccifera, C. crispata,
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20 BRIAN JOHN COPPINS Micarea The g
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22 BRIAN JOHN COPPINS Morphology Th
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24 BRIAN JOHN COPPINS (ii) Goniocys
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26 BRIAN JOHN COPPINS A B Fig. 2 'M
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28 BRIAN JOHN COPPINS 3 00
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30 BRIAN JOHN COPPINS Fig. 5 Sketch
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32 BRIAN JOHN COPPINS Fig. 6 Ascus
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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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42 BRIAN JOHN COPPINS Fig. 16 M. in
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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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62 BRIAN JOHN COPPINS Fig. 34 Two t
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64 BRIAN JOHN COPPINS Table 1 Conid
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66 BRIAN JOHN COPPINS Fig. 36 Sugge
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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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84 BRIAN JOHN COPPINS In the case o
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86 BRIAN JOHN COPPINS 6 - 5h 4 3 2
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. 88 BRIAN JOHN COPPINS the epithec
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90 BRIAN JOHN COPPINS fM. elachista
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92 BRIAN JOHN COPPINS Saxicolous ha
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LICHEN GENUS MICAREA IN EUROPE 95 T
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LICHEN GENUS MICAREA IN EUROPE 97 a
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Infrageneric considerations LICHEN
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LICHEN GENUS MICAREA IN EUROPE 201
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conferanda (Lecidea) 205 confusula
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populina (Lecidea, Micarea) 31,32 p
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