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Bulletin of the British Museum (Natural History)

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152 BRIAN JOHN COPPINS<br />

c. 40-60 /Ltm diam, semi-immersed to sessile, walls green, K—;conidia (microconidia) fusiform-<br />

cylindrical, straight, 4-8-7x0-8-l /itm.<br />

Chemistry: Thallus K— , C+<br />

red or C— , PD; sections <strong>of</strong> apo<strong>the</strong>cia C— ; t.l.c: gyrophoric acid<br />

(healthy thalli), or no substances or gyrophoric acid in trace amounts (poorly developed or<br />

disrupted thalH).<br />

Observations: Micarea melaena is one <strong>of</strong> <strong>the</strong> commonest species <strong>of</strong> <strong>the</strong> genus, especially in<br />

nor<strong>the</strong>rn Europe, and it displays a considerable amount <strong>of</strong> variation in thallus appearance and<br />

internal apo<strong>the</strong>cial pigmentation. However, my examinations <strong>of</strong> over 200 specimens have<br />

revealed many intermediates between <strong>the</strong> known extremes. When well-developed, <strong>the</strong> thallus is<br />

composed <strong>of</strong> cohering, small, granular areolae and is pale buff to dull green in colour. In<br />

exposed situations <strong>the</strong> thallus <strong>of</strong>ten becomes a dark green-black due to <strong>the</strong> more intense<br />

pigmentation <strong>of</strong> <strong>the</strong> outermost hyphae <strong>of</strong> <strong>the</strong> granules. The appearance <strong>of</strong> <strong>the</strong> thallus can also be<br />

affected by <strong>the</strong> presence <strong>of</strong> o<strong>the</strong>r organisms (see description and p. 00). When on lignum, in dry<br />

exposed situations <strong>the</strong> thallus may become much reduced to a fine scattering <strong>of</strong> tiny (less than 30<br />

/xm diam), blackish granules amongst <strong>the</strong> wood fibres. Such forms proved to be very common,<br />

during my explorations in Sweden.<br />

M. melaena also exhibits much variation with regard to <strong>the</strong> colouration <strong>of</strong> its apo<strong>the</strong>cial tissues<br />

(especially <strong>the</strong> hypo<strong>the</strong>cium). This is due to <strong>the</strong> presence <strong>of</strong> three (probably biosyn<strong>the</strong>tically<br />

related) pigments, varying in <strong>the</strong>ir relative amounts and distribution. These pigments, all <strong>of</strong><br />

which react HNO3+ purple-red, can be roughly identified as pigment A: green, K-l- green<br />

intensifying; pigment B: purple, K+ green; pigment C: purple, K+ purple intensifying.<br />

In <strong>the</strong> epi<strong>the</strong>cium and hymenium pigment A is <strong>of</strong>ten found alone, but sometimes it is replaced<br />

in <strong>the</strong> lower hymenium (or in extreme cases throughout <strong>the</strong> entire hymenium and epi<strong>the</strong>cium)<br />

by pigment C. Pigment C is usually dominant in <strong>the</strong> hypo<strong>the</strong>cium, but is <strong>of</strong>ten replaced in <strong>the</strong><br />

upper and outer parts (or more rarely ± throughout) by pigment B. If thin sections <strong>of</strong> apo<strong>the</strong>cia<br />

in which pigment B is dominant are mounted in K <strong>the</strong> lower central ('core') <strong>of</strong> <strong>the</strong> hypo<strong>the</strong>cium<br />

retains a purplish tinge (<strong>the</strong> rest having turned green), indicating <strong>the</strong> presence <strong>of</strong> a small amount<br />

<strong>of</strong> pigment C. The same type <strong>of</strong> variation, presumably involving identical (or very similar)<br />

pigments, is found in several o<strong>the</strong>r species, e.g. M. crassipes and M. sylvicola.<br />

The pycnidia <strong>of</strong> M. melaena are very inconspicuous and apparently sparingly produced.<br />

However, a close scrutiny <strong>of</strong> about 100 specimens revealed ten collections with <strong>the</strong> microconi-<br />

dial state and only two (e.g. Coppins 6041) with <strong>the</strong> macroconidial state. A mesoconidial state<br />

was not found but may well occur.<br />

Habitat and distribution: M. melaena is widely distributed and especially common in northwest<br />

Europe. Although, for a Micarea, it is a conspicuous species, it is <strong>of</strong>ten overlooked as a<br />

black or charred stain on <strong>the</strong> tops <strong>of</strong> stumps and peaty turf! It is a common inhabitant <strong>of</strong> stumps<br />

and fallen decorticate trunks in upland woodlands, or in lowland woodlands on very acid,<br />

nutrient-poor soils, and can exist in shaded to extremely exposed, sun-baked situations. In<br />

upland woods it can be found on <strong>the</strong> corticate trunks <strong>of</strong> healthy trees (especially Betula,<br />

Quercus, and Pinus). In areas heavily polluted by SO2 and its derivatives (e.g. West Yorkshire<br />

conurbation) it has been found on <strong>the</strong> acidified bark <strong>of</strong> Acer and Ulmus. Occurrences on<br />

decaying timber work (e.g. fence posts, gate rails and shingles) are not uncommon, especially in<br />

upland districts. Lichens associated with M. melaena on lignum and bark in <strong>the</strong> <strong>British</strong> Isles<br />

include Bryoria capillaris, B. fuscescens, Cladonia spp., Hypocenomyce scalaris, Hypogymnia<br />

physodes, Lecidea granulosa agg., L. icmalea, Micarea denigrata, M. prasina, Mycoblastus<br />

sterilis, Ochrolechia turneri, Parmelia saxatilis, Parmeliopsis spp., Platismatia glauca, Trapelia<br />

corticola, and Usnea spp.<br />

M. melaena is <strong>of</strong>ten terricolous, growing on oligotrophic, peaty soils, in montane regions or<br />

lowland heaths. In <strong>the</strong> English lowlands it is recorded from heaths in <strong>the</strong> Sussex Weald and east<br />

Norfolk. It is common on ± bare peat in <strong>the</strong> Pennines and Scottish highlands, but in <strong>the</strong> latter it<br />

is usually replaced near <strong>the</strong> higher summits (over c. 1000 m) by M. turfosa. Associated lichens on<br />

peaty turf in Britain include 'Botrydina' (i.e. Omphalina spp.), Cladonia coccifera, C. crispata,

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