Bulletin of the British Museum (Natural History)
Bulletin of the British Museum (Natural History)
Bulletin of the British Museum (Natural History)
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LICHEN GENUS MICAREA IN EUROPE 129<br />
forms), sometimes emergent, hyaline grey or black; walls hyaline but usually dilute olivaceous<br />
or brownish, and K+ violet in <strong>the</strong> upper (exposed) parts; in endoxylic forms, walls olivaceous<br />
(K+ violet) throughout; <strong>of</strong> three types: (a) 60-150 ^im diam, ostioles eventually gaping; conidia<br />
(macroconidia) , curved or hamate, (l-)3-septate, 12-24xc. 1 /xm; (b) 80-160 ^im diam, ostioles<br />
<strong>of</strong>ten gaping: conidia (mesoconidia) short cylindric or obovate, sometimes faintly biguttulate,<br />
2-8^-5(-5)xl-2-l-8 fxm <strong>of</strong>ten extruding from <strong>the</strong> ostioles as a conspicuous white blobs; (c)<br />
30-50 /xm, ostioles not gaping; conidia (microconidia) narrowly fusiform or bacilliform,<br />
(4-5)5-7-5x0-7-0-8/xm.<br />
Chemistry: Thallus K— , PD— ; sections <strong>of</strong> thallus and apo<strong>the</strong>cia C+ orange-red, rarely C—<br />
(also, parts with dull olivaceous pigment, C+ violet). If thallus is heavily parasitised and scurfy<br />
<strong>the</strong> C+ orange-red reaction may be difficult to obtain and gyrophoric acid may not be detectable<br />
by t. I.e.<br />
Observations: Micarea denigrata is a common and extremely polymorphic species exhibiting a<br />
wide range <strong>of</strong> genotypic and phenotypic variation. Its thallus can vary from being completely<br />
endoxylic to forming a thick, granular-verrucose, areolate crust. The colour <strong>of</strong> <strong>the</strong> apo<strong>the</strong>cia<br />
varies from pallid, through grey or brown, to black in a ± direct relationship with exposure to<br />
Ught. Specimens may be abundantly fertile with numerous, large, mature apo<strong>the</strong>cia. On <strong>the</strong><br />
o<strong>the</strong>r hand, <strong>the</strong>y may be ± sterile with just a few, scattered immature apo<strong>the</strong>cia (or even none at<br />
all) but with numerous pycnidia containing mesoconidia; alternatively <strong>the</strong> latter may be<br />
replaced by pycnidia containing microconidia and, or, macroconidia. Indeed, all possible<br />
combinations <strong>of</strong> anamorphic states have been found and a few collections (e.g. Coppins 1888<br />
and 8384) have all three states, plus apo<strong>the</strong>cia, on <strong>the</strong> same thallus. Spore length is usually in <strong>the</strong><br />
range <strong>of</strong> 9-14 /^tm, but extreme forms are known in which <strong>the</strong> range is 8-10 /x,m, or 12-18 ^m.<br />
Gyrophoric acid is usually present in <strong>the</strong> thallus and/or <strong>the</strong> apo<strong>the</strong>cia (and easily demonstrated<br />
by tests with C), but in some specimens its concentration is low and only detectable with<br />
certainty by t.l.c. ; in a few diminutive, endoxyhc forms I have been unable to detect gyrophoric<br />
ei<strong>the</strong>r by a C test, or by t.l.c. In short, M. denigrata exhibits extreme variation. Despite this, with<br />
care and patience <strong>the</strong> species is rarely difficult to separate from o<strong>the</strong>r, similar species. The<br />
biggest problems I have experienced are where <strong>the</strong> decision has to be made between what could<br />
be a diminutive form <strong>of</strong> M. denigrata with small spores, or a form oiM. misella with an unusually<br />
high proportion <strong>of</strong> 1 -septate spores and without its characteristic stalked pycnidia (that contain<br />
mesoconidia). Such decisions are settled by <strong>the</strong> careful observation and measurement <strong>of</strong><br />
paraphyses, and microconidia (see couplet 11 <strong>of</strong> <strong>the</strong> main key and Table 7). Shade forms <strong>of</strong> M.<br />
denigrata with brown, ± globose apo<strong>the</strong>cia have been confused with M. elachista, but <strong>the</strong> latter<br />
has a brown (K+ dissolving) epi<strong>the</strong>cial pigment, a more complex thallus structure (with a<br />
'cortex' and amorphous covering layer), distinctive pycnidia, and always lacks gyrophoric acid<br />
(sections <strong>of</strong> thallus and apo<strong>the</strong>cia C— ). Forms <strong>of</strong> M. denigrata with a scurfy-granular thallus can<br />
be similar to M. prasina, but microscopic examination will show <strong>the</strong> thallus appearance to be<br />
caused by its disruption by invading foreign fungi and algae, and not by <strong>the</strong> presence <strong>of</strong> <strong>the</strong> small<br />
± discrete granules (goniocysts) characteristic <strong>of</strong> M. prasina. Fur<strong>the</strong>rmore, M. prasina never<br />
gives C+ orange-red (gyrophoric acid) reactions, and it usually has broader, rarely curved<br />
spores with more rounded apices. The closest relative to M. denigrata is M. nitschkeana, which is<br />
± identical with regard to thallus structure, pycnidial types, apo<strong>the</strong>cia structure, pigmentation,<br />
and chemistry. Morphologically <strong>the</strong> only important difference is that <strong>the</strong> mature spores <strong>of</strong> M.<br />
nitschkeana are 3-septate and more consistently curved (cf. Figs 13 and 24B). In addition, <strong>the</strong><br />
two species differ in <strong>the</strong>ir preferance <strong>of</strong> substrata: M. denigrata favouring lignum or dead bark <strong>of</strong><br />
old tree trunks or stumps, and M. nitschkeana favouring corticate twigs and small branches <strong>of</strong><br />
trees and shrubs. However, <strong>the</strong>re is some degree <strong>of</strong> overlap as M. nitschkeana is occasionally<br />
found on <strong>the</strong> lignum <strong>of</strong> fence posts, but, on <strong>the</strong> o<strong>the</strong>r hand, I have never encountered M.<br />
denigrata on attached living, corticate twigs. In <strong>the</strong> field, well developed specimens <strong>of</strong> M.<br />
denigrata sometimes have a superficial resemblance to M. cinerea, M. lignaria, and M.<br />
peliocarpa, but <strong>the</strong>se three species have K— hymenia and larger, 3 (or more)-septate spores.<br />
Diminutive endoxyhc forms <strong>of</strong> M. denigrata are indistinguishable in <strong>the</strong> field from <strong>the</strong> several,