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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O' M'<br />
<strong>Bulletin</strong> <strong>of</strong> <strong>the</strong><br />
<strong>British</strong> <strong>Museum</strong> (<strong>Natural</strong> <strong>History</strong>)<br />
A taxonomic study <strong>of</strong> <strong>the</strong> lichen genus<br />
Micarea in Europe<br />
Brian John Coppins<br />
Botany series Vol 11 No 2 29 September 1983
A taxonomic study <strong>of</strong> <strong>the</strong> lichen genus Micarea in<br />
Europe<br />
Brian John Coppins ](i,<br />
Royal Botanic Garden, Inverleith Row, Edinburgh, EH3 5LR<br />
Contents<br />
Synopsis 18<br />
Historical background 19<br />
Lecideaceae 19<br />
Micarea 20<br />
Materials 20<br />
Methods 21<br />
Field studies 21<br />
Light microscopy 21<br />
Thin-layer chromatography 21<br />
Nomenclature 21<br />
Morphology 22<br />
Thallus 22<br />
Phycobionts 25<br />
Cephalodia 25<br />
Apo<strong>the</strong>cia: external features 26<br />
Apo<strong>the</strong>cia: internal features 30<br />
Anamorphs (conidial states) 63<br />
Chemistry 84<br />
Lichen substances 84<br />
Pigments 87<br />
Ecology 88<br />
General habitats 88<br />
Specific habitats 90<br />
Distribution 93<br />
Britain 93<br />
Europe 94<br />
World 96<br />
The genus Micarea 96<br />
Intergeneric considerations 97<br />
Suprageneric considerations 98<br />
Infrageneric considerations 99<br />
Keys to species 100<br />
Guide to keys & identifications 100<br />
Key to European species 101<br />
Key to European species that may occur without apo<strong>the</strong>cia 107<br />
The species 108<br />
1. M. adnata 108<br />
2. M. alabastrites HO<br />
3. M. anterior 112<br />
4. M. assimilata 114<br />
5<br />
.<br />
M. bauschiana<br />
6. M. botryoides 118<br />
7. M. cinerea 121<br />
8. M.contexta • 124<br />
9. M. crassipes 125<br />
10. M.curvata 126<br />
Bull. Br. Mas. nat. Hist. (Bot.) 11 (2): 17-214<br />
~<br />
W. u-^^<br />
H^<br />
Issued 29 September 1983
.<br />
18 BRIAN JOHN COPPINS<br />
11. M.denigrata 127<br />
12. M. elachista 131<br />
13. M. eximia 134<br />
14. M. globulosella 134<br />
15. M.hedlundii 135<br />
16. M. incrassata 137<br />
17. M. intrusa 138<br />
18. M.leprosula 140<br />
19. M.lignaria 142<br />
20. M.lithinella 147<br />
21. M.lutulata 148<br />
22. M. melaena 150<br />
23. M. melaenida 154<br />
24. M. melaeniza 155<br />
25. M. melanobola 156<br />
26. M.misella 158<br />
27. M. muhrii 160<br />
28. M. myriocarpa 161<br />
29. M.nigella 163<br />
30. M. nitschkeana 165<br />
31 M. olivacea 167<br />
32. M. osloensis 169<br />
33. M. peliocarpa 169<br />
34. M. prasina 173<br />
35. M. pycnidiophora 179<br />
36. M. rhabdogena 181<br />
37. M.stipitata 182<br />
38. M. subleprosula 182<br />
39. M.subnigrata 183<br />
40. M. subviolascens 185<br />
41 M. sylvicola 186<br />
42. M. syno<strong>the</strong>oides 188<br />
43. M. ternaria 190<br />
44 M. tuberculata 192<br />
45. M.turfosa 194<br />
Excluded taxa 196<br />
Index to exsiccatae 203<br />
Acknowledgements 206<br />
References 207<br />
Index 210<br />
Synopsis<br />
A taxonomic revision is presented for <strong>the</strong> lichen genus Micarea in Europe, with special emphasis on those<br />
species occurring in <strong>the</strong> <strong>British</strong> Isles. In brief, this genus is here circumscribed to include most crustose<br />
Hchens with lecideine (biatorine) apo<strong>the</strong>cia, a poorly defined excipulum (sometimes absent altoge<strong>the</strong>r) <strong>of</strong><br />
radiating paraphysis-like hyphae, a non-granular epi<strong>the</strong>cium, Lecanora-type, 8-spored asci, simple to<br />
multiseptate, hyaline ascospores, and a 'grass-green' phycobiont usually <strong>of</strong> <strong>the</strong> so-called 'micareoid' type.<br />
Comparisons and possible relationships with similar genera are discussed. Noteworthy discoveries made<br />
during this study include <strong>the</strong> finding <strong>of</strong> cephalodia in three species, dimorphic paraphyses in several<br />
species, and a wide array <strong>of</strong> anamorphic forms, with three species each found to have three conidial states.<br />
Type studies have been made for nearly all names known, as well as those suspected, to be referable to<br />
Micarea in its present, wider concept. Forty-five species are recognised, <strong>of</strong> which 31 are confirmed from<br />
Britain. Seven species are new to science: Micarea adnata, M. curvata, M. hedlundii, M. muhrii, M. nigella,<br />
M. olivacea, and M. myriocarpa Vezda & V. Wirth ex Coppins. Several described species are included<br />
in Micarea for <strong>the</strong> first time, and additional name changes are required for nomenclatural reasons; nine<br />
new combinations result: M. assimilata (Nyl.), M. crassipes (Th.Fr.), M. elachista (Korber) Coppins «fe<br />
R. Sant. , M. globulosella (Nyl) , M. intrusa (Th.Fr.) Coppins & Kilias, M. lignaria war. endoleuca (Leigh ton)<br />
M. melaenida (Nyl.), M. melanobola (Nyl.), and M. subviolascens (Magnusson). Several taxa are ex-<br />
,
LICHEN GENUS MICAREA IN EUROPE 19<br />
eluded from <strong>the</strong> genus and <strong>the</strong> new combinations, Psilolechia clavulifera (Nyl.) and Bacidia prasinata<br />
(Tuck.), are proposed. Keys for <strong>the</strong> identification <strong>of</strong> all <strong>the</strong> accepted European taxa are given. The taxonomic<br />
parts are preceded by an outline <strong>of</strong> <strong>the</strong> historical background to <strong>the</strong> study <strong>of</strong> Micarea, and details <strong>of</strong><br />
materials and methods employed in this study. Detailed accounts <strong>of</strong> <strong>the</strong> morphology, chemistry, and<br />
ecology in <strong>the</strong> genus are provided , and a discussion <strong>of</strong> distributions is supported by maps for <strong>the</strong> <strong>British</strong> taxa.<br />
All Micarea species occur on acidic, nutrient poor substrata, and most are confined to cool-temperate,<br />
boreal, or oceanic regions; a few occur in arctic-alpine areas but <strong>the</strong> genus is poorly represented in<br />
dry, lowland, Mediterranean regions. Prior to this study, most <strong>of</strong> <strong>the</strong> species were little-known or<br />
misunderstood; clarification <strong>of</strong> <strong>the</strong>ir taxonomy has been achieved by paying particular attention to <strong>the</strong>ir<br />
anamorphic states, chemistry (including pigmentation), and detailed anatomy. Consideration <strong>of</strong> <strong>the</strong><br />
distribution and ecology <strong>of</strong> <strong>the</strong> species has proved invaluable in ordering <strong>the</strong> taxonomic chaos which previously<br />
surrounded <strong>the</strong> notoriously variable species <strong>of</strong> <strong>the</strong> genus.<br />
Lecideaceae<br />
Historical background<br />
Until recently <strong>the</strong> circumscription <strong>of</strong> <strong>the</strong> Lecideaceae (and <strong>of</strong> <strong>the</strong> genera within it) had changed<br />
httle from that adopted by Zahlbruckner (1926). It included most Uchens with <strong>the</strong> following<br />
combination <strong>of</strong> characters: a crustose to squamulose thallus, a 'grass-green' phycobiont<br />
(excluding Trentepohlia and Phycopeltis) , ± immersed to sessile, disc-like apo<strong>the</strong>cia without a<br />
thalhne margin, mainly colourless spores, and an absence <strong>of</strong> parietin (or related pigments) and<br />
(or) polarilocular spores. The principal genera in <strong>the</strong> family (e.g. Lecidea, Catillaria, Bacidia,<br />
Biatorella, Mycoblastus, Lopadium, Bombyliospora, and Toninia) were separated mainly on<br />
<strong>the</strong> basis <strong>of</strong> spore characters, i.e. size, septation, and number per ascus. This classification<br />
largely ignored many features which (according to modern mycological concepts) now merit<br />
careful consideration, although <strong>the</strong>y were used to varying degrees for <strong>the</strong> delimitation <strong>of</strong> species<br />
or infrageneric categories above this rank. In brief, <strong>the</strong>se features involve <strong>the</strong> structure <strong>of</strong> asci,<br />
excipular and hypo<strong>the</strong>cial tissues, paraphyses and anamorphs, ontogeny, finer aspects <strong>of</strong> thallus<br />
structure, and nature and location <strong>of</strong> pigments and lichen substances. In addition, investigations<br />
<strong>of</strong> <strong>the</strong> phycobiont(s) and considerations <strong>of</strong> ecology and phytogeography <strong>of</strong>ten provide valuable<br />
supplementary information. However, <strong>the</strong> use <strong>of</strong> some <strong>of</strong> <strong>the</strong>se features in an attempt to define<br />
more natural genera is not a purely recent phenomenon. Several lichenologists working in <strong>the</strong><br />
1850s and 1860s made bold attempts in this direction. With regard to <strong>the</strong> Lecideaceae s. lat., two<br />
lichenologists deserving special mention are G. W. Korber (who introduced Lecidella, Lopadium,<br />
Pyrrhospora, Schaereria, Schadonia, and Steinia) and A. B. Massalongo (who introduced<br />
Catillaria, Psilolechia, Sarcosagium, Scoliciosporum, Strangospora, and Toninia). In <strong>the</strong> latter<br />
half <strong>of</strong> <strong>the</strong> 19th century lichenology came under <strong>the</strong> almost monarchical influence <strong>of</strong> William<br />
Nylander, whose simplistic generic concepts gained precedence over <strong>the</strong> more far-sighted works<br />
<strong>of</strong> Korber, Massalongo, and o<strong>the</strong>rs. From <strong>the</strong> 1870s right up to <strong>the</strong> 1950s <strong>the</strong>re were few<br />
attempts to reassess <strong>the</strong> generic concepts <strong>of</strong> Nylander or <strong>the</strong> slightly more complex, but no less<br />
artificial, system <strong>of</strong> Zahlbruckner. Between about 1929 and 1954 <strong>the</strong> French lichenologist M. G.<br />
B. Choisy resurrected many <strong>of</strong> <strong>the</strong> old and more or less forgotten genera, and created several<br />
new ones (e.g. Haplocarpon [= Huilia], Hypocenomyce, Trapelia, and Tremolecia). Unfortu-<br />
nately, Choisy's works made little impact at <strong>the</strong> time and it was not until <strong>the</strong> mid-1960s that<br />
lichenologists began to look more carefully at <strong>the</strong> delimitation <strong>of</strong> genera. Recent investigations<br />
have led to <strong>the</strong> reinstatement (although <strong>of</strong>ten with emendations) <strong>of</strong> many <strong>of</strong> <strong>the</strong>se genera and<br />
many new genera have had to be described (e.g. Fuscidea, Herteliana, Melanolecia, Trapeliop-<br />
sis, Tylothallia, and Vezdaea). Most are included in <strong>the</strong> key to European lichen genera in Poelt<br />
& Vezda (1981). Despite <strong>the</strong> many advances made during <strong>the</strong> last 15 years, it will be several<br />
decades before a reasonably natural generic classification within <strong>the</strong> Lecideaceae s. lat. is<br />
achieved. The enormity <strong>of</strong> <strong>the</strong> task can be appreciated from <strong>the</strong> fact that Zahlbruckner<br />
(1921-40) accepted no less than 1450 species in <strong>the</strong> genus Lecidea alone! In addition to <strong>the</strong> high<br />
number <strong>of</strong> taxa involved , fur<strong>the</strong>r problems arise from <strong>the</strong> locating and obtaining on loan suitable<br />
(including type) material, and <strong>the</strong> many difficulties in observing and interpreting many<br />
microscopical, morphological, and ontogenetic features.
20 BRIAN JOHN COPPINS<br />
Micarea<br />
The genus Micarea was first validly described in 1825 by Elias Fries in his Systerna orbis<br />
vegetabilis (see p. 96), and was placed in his 'Tribus Collemaceae' on account <strong>of</strong> its ra<strong>the</strong>r<br />
gelatinous thallus, although he noted that it had Lecidea-hke apo<strong>the</strong>cia. The generic name was<br />
little used by most 19th century lichenologists, although it was accepted with <strong>the</strong> single species<br />
{M. prasina) by a few such as Korber (1855). Towards <strong>the</strong> end <strong>of</strong> that century, J. T. Hedlund<br />
submitted his doctoral <strong>the</strong>sis to <strong>the</strong> University <strong>of</strong> Uppsala. In this work (Hedlund, 1892) he<br />
adopted and emended <strong>the</strong> genus to include 20 species, and his circumscription <strong>of</strong> <strong>the</strong> genus is<br />
essentially <strong>the</strong> same as that accepted by Vezda & Wirth (1976) and myself, although many<br />
species have since been added. Hedlund's sagacious work was evidently too revolutionary for his<br />
time, and it did not achieve international recognition. It seems that Hedlund was disillusioned<br />
and had difficulties in finding a position. He turned to horticulture (especially dendrology) and<br />
<strong>the</strong>n became an authority on Sorbus. Although Hedlund's obvious talents were not lost to<br />
botany, <strong>the</strong>y were sadly lost to lichenology, especially <strong>the</strong> study <strong>of</strong> microlichens.<br />
Vezda & Wirth (1976) slightly expanded Hedlund's concept <strong>of</strong> Micarea by including Lecidea<br />
sylvicola and related species (all without 'micareoid' algae), and also Bacidia beckhausii. Apart<br />
from <strong>the</strong> exclusion <strong>of</strong> B. beckhausii, my own concepts are much <strong>the</strong> same, although I have<br />
emended <strong>the</strong> genus very slightly so as to include species such as Lecidea (Helocarpon) crassipes<br />
and Catillaria intrusa.<br />
The first species <strong>of</strong> Micarea to be described was Lecidea [Micarea] lignaria Ach. (1808). By<br />
1850 only seven <strong>of</strong> <strong>the</strong> currently accepted 45 European species were validly published. At about<br />
this time lichenologists began to make use <strong>of</strong> better quality microscopes and <strong>the</strong> additional<br />
characters <strong>the</strong>y revealed. By <strong>the</strong> end <strong>of</strong> <strong>the</strong> nineteenth century 34 <strong>of</strong> <strong>the</strong> accepted species had<br />
been described. The European Micarea flora was increased by only one species (Lecidea<br />
subviolascens) between 1900 and 1960. In 1961 Vezda published Bacidia [Micarea] subleprosula,<br />
and as a result <strong>of</strong> <strong>the</strong> present studies (some in collaboration with Mr P. W. James, Dr A.<br />
Vezda, and Dr V. Wirth) a fur<strong>the</strong>r nine species have been described. In <strong>the</strong> future a few<br />
additional species will probably be added to <strong>the</strong> European flora, but <strong>the</strong> greatest expansion<br />
within <strong>the</strong> genus will come from <strong>the</strong> study <strong>of</strong> extra-European collections; indeed, many<br />
European, undescribed, and described (in o<strong>the</strong>r genera) species are already known to me from<br />
parts <strong>of</strong> <strong>the</strong> world outside Europe.<br />
Materials<br />
During <strong>the</strong> course <strong>of</strong> this study I have examined about 3,000 specimens (c. 1800 from <strong>the</strong> <strong>British</strong><br />
Isles) attributable to Micarea, plus a fur<strong>the</strong>r c. 500 specimens which have been referred to o<strong>the</strong>r<br />
genera. Material has been received on loan from (or studied in) <strong>the</strong> following institutional<br />
herbaria: ABD, ANGUC, BEL, BERN, BG, BM, BON, C, DBN, DEE, DUKE, E, G,<br />
GLAM, GZU, H, HAMU, HBG, HEX, IMI, K (now in BM; cited as BM ex K), L, LD, LIV,<br />
LSR, M, MANCH, NMW, NWH, O, S, STD, STU, SUN, TUR, U, UPS, VER, WCR, WIS,<br />
WRSL; abbreviations according to Holmgren et al. (1981). In addition, numerous specimens<br />
have been received on loan from private herbaria. From <strong>the</strong> <strong>British</strong> Isles <strong>the</strong>se are: Dr H. J. M.<br />
Bowen (Oxford), Dr R. W. M. Corner (Penrith), Mr I. P. Day (Carlisle), Dr U. K. Duncan<br />
(Arbroath; lichen herbarium recently gifted to E), Dr A. Fletcher (Leicester), Mr V. J.<br />
Giavarini (Parkstone, Dorset), Dr O. L. Gilbert (Sheffield), Mr R. Gomm (Taunton), Rev. G.<br />
G. Graham (Hunwick, Co. Durham), Mr A. Henderson (Leeds), Dr C. J. B. Hitch (Saxmundham,<br />
Suffolk), Dr P. D. Hulme (Aberdeen), Dr A. R. Pentecost (Royal Tunbridge Wells), Dr<br />
F. Rose (Liss, Hampshire; many specimens now in BM), Dr M. R. D. Seaward (Bradford), Mr<br />
J. F. Skinner (Sou<strong>the</strong>nd-on-Sea), Dr P. B. Topham (Dundee), Mr R. G. Woods (Newbridge on<br />
Wye, Powys); and from elsewhere in Europe: Dr J. Hafellner, DrH. Mayrh<strong>of</strong>er, and Pr<strong>of</strong>. Dr J.<br />
Poelt (all Graz, Austria), Mr L.-E. Muhr (Karlskoga, Sweden), Dr A. Vezda (Brno, Czechoslovakia),<br />
and Dr V. Wirth (Ludwigsburg, W. Germany).<br />
When studying a group <strong>of</strong> much misunderstood lichens it is a rewarding exercise to investigate<br />
folders <strong>of</strong> o<strong>the</strong>r superficially similar (but strictly unrelated) taxa, especially those that occur in
LICHEN GENUS MICAREA IN EUROPE 21<br />
similar habitats. During this study numerous specimens <strong>of</strong> Micarea have been found in folders <strong>of</strong><br />
widely misinterpreted ('dustbin') names, such as ' Bacidia sphaeroides' , 'Catillaria erysiboides'<br />
and Lecidea vemails'<br />
.<br />
Copies <strong>of</strong> <strong>the</strong> list <strong>of</strong> specimens examined have been lodged at BM, DBN, E, GZU, M, NMW,<br />
UPS and US.<br />
Field studies<br />
Methods<br />
I have attempted to observe as many as possible <strong>of</strong> <strong>the</strong> accepted species in <strong>the</strong>ir natural habitats<br />
because such experience is invaluable for <strong>the</strong> appreciation <strong>of</strong> environmentally controlled<br />
variation and ecological requirements. For this end I have made field studies in most parts <strong>of</strong><br />
mainland Britain, and also S.E. Ireland, Denmark (N. Jylland), mid- and N. Sweden. In<br />
addition, I have collected Micarea specimens during earlier expeditions to France (Bretagne)<br />
and western Ireland. I have been successful in finding two-thirds <strong>of</strong> <strong>the</strong> European species, and all<br />
but two {M. asslmllata and M. subleprosula) <strong>of</strong> <strong>the</strong> 31 <strong>British</strong> species.<br />
Light microscopy<br />
Observations and measurements <strong>of</strong> external features were mostly made at x50 using a Vickers<br />
stereomicroscope equipped with a measuring eyepiece. Internal features were investigated with<br />
a Wild M20 microscope which was fitted with a drawing tube and a 2-5 x adapter for <strong>the</strong> drawing<br />
<strong>of</strong> spores and conidia etc. Most sections were made by hand with a razor blade, but some were<br />
cut by a freezing microtome. Sections were usually mounted in water, followed by (or directly<br />
in) 10% or 50% KOH(K), domestic bleach(C), 50% HNO3, or ammoniacal erythrosin (0-5g<br />
erythrosin in 100 ml 10% ammonia solution), but o<strong>the</strong>r mountants such as cotton-blue in<br />
lactophenol (LCB), congo red, and alcian blue were also used at times. Tests for amyloid<br />
reactions were made by mounting directly in Lugol's iodine solution (Ig iodine and 2g potassium<br />
iodide in 300 ml distilled water), or in this solution following treatment with 10% KOH. More<br />
permanent preparations were made by ringing mounts in LCB with nail varnish, or by mounting<br />
in polyvinyl alcohol (PVA) or cotton blue in PVA (see Omar etal, 1979). For fur<strong>the</strong>r details on<br />
techniques see also 'Guide to keys and identifications' (p. 100).<br />
Thin-layer chromatography<br />
The t.l.c. techniques employed were those described by Walker & James (1980) which are based<br />
on <strong>the</strong> standard method <strong>of</strong> Culberson (1972) . For <strong>the</strong> purposes <strong>of</strong> routine analysis only two <strong>of</strong> <strong>the</strong><br />
three basic solvent systems were found to be necessary (i.e. H.E.F. and T.D.A.).<br />
Nomenclature<br />
In <strong>the</strong> list <strong>of</strong> synonyms for a given species each entry begins with <strong>the</strong> oldest valid name<br />
(basionym) and is followed with later combinations included in Hedlund (1892), Smith (1911,<br />
1926), and James (1965^), and some o<strong>the</strong>rs which have important nomenclatural implications.<br />
Zahlbruckner (1921-40) and Lamb (1963) should be consulted for additional combinations. The<br />
entries are listed chronologically, except for entirely invalid (or illegitimate) and misapplied<br />
names which are included at <strong>the</strong> end <strong>of</strong> <strong>the</strong> lists.<br />
Abbreviations <strong>of</strong> authors are according to Hawksworth (1980); those <strong>of</strong> journals to <strong>the</strong> third<br />
[1980] edition <strong>of</strong> Serial Publications In <strong>the</strong> <strong>British</strong> <strong>Museum</strong> (<strong>Natural</strong> <strong>History</strong>) Library; and those<br />
<strong>of</strong> books to Hawksworth (1974) or Stafleu & Cowan (1976-81).<br />
Apart from those newly described in recent years, few names in Micarea have been formally<br />
typified (according to Art. 7 <strong>of</strong> <strong>the</strong> Code); all cited lectotypes and neotypes are selected in this<br />
work, unless o<strong>the</strong>rwise indicated.<br />
The nomenclature <strong>of</strong> lichens not treated in detail mainly follows Hawksworth et al. (1980) , but<br />
a few later changes are used.<br />
,
22 BRIAN JOHN COPPINS<br />
Morphology<br />
Thallus<br />
Thallus structure in Micarea is crustose and basically simple, but never<strong>the</strong>less, encompasses<br />
much interspecific and intraspecific variation. For <strong>the</strong> purposes <strong>of</strong> explanation thallus morphology<br />
can be roughly divided into five types. [Note: <strong>the</strong> discussion <strong>of</strong> <strong>the</strong> first type includes some<br />
general features applicable to all types.]<br />
(i) Areolate-type (Fig. lA-C)<br />
The term areolae (pi.) is used here to describe discrete, rounded (when viewed from above),<br />
flattened, or more usually convex (sometimes ± globose) portions <strong>of</strong> <strong>the</strong> thallus which have<br />
developed directly from <strong>the</strong> prothallus lying in or on <strong>the</strong> substratum. These should be<br />
distinguished from <strong>the</strong> (<strong>of</strong>ten angular) segments derived from <strong>the</strong> cracking <strong>of</strong> a previously<br />
continuous crust. The areolae in Micarea mostly range from <strong>the</strong> 0-06 to 0.2 mm diam, but are<br />
characteristically larger in certain species. Thalli comprising smaller, discrete, granular, soredia-<br />
Uke structures (goniocysts) are dealt with in <strong>the</strong> next category (ii).<br />
Species that have well-developed, convex areolae sometimes appear ± squamulose, e.g. M.<br />
assimilata, M. incrassata, M. lignaria (as in type <strong>of</strong> Lecidea trisepta var. polytropoides) , M.<br />
melaenida, and M. subviolascens. Catillaria zsakii (a synonym <strong>of</strong> M. melaenida) was placed in<br />
<strong>the</strong> squamulose genus Toninia Massal. by Lettau, but its thallus is not truly squamulose and it<br />
was excluded from Toninia by Baumgartner (1979). In M. subviolascens (and to a lesser extent<br />
in some o<strong>the</strong>r species) <strong>the</strong> areolae become confluent and <strong>the</strong> resultant crust becomes secondarily<br />
cracked into 'islands' (each containing several primary areolae), thus giving <strong>the</strong> thallus a ra<strong>the</strong>r<br />
squamulose appearance. The areolae in most species are whitish or pale to medium grey, and<br />
<strong>of</strong>ten tinged dull greenish, bluish or brownish. The areolae <strong>of</strong> M. lignaria var. endoleuca are<br />
characterically ivory or cream-yellow. More vivid yellow, citrine or orange thallus colourations<br />
are not known in Micarea.<br />
A few whitish, arachnoid, prothalline hyphae are sometimes visible in specimens with<br />
scattered areolae, but a thick prothallus as found, for example, in some species <strong>of</strong> Phyllopsora<br />
(Swinscow & Krog, 1981) is unknown in Micarea. The thallus in most Micarea spp. is effuse and<br />
wide-spreading, and even when thalli form small patches amongst o<strong>the</strong>r lichens, a delimiting<br />
hypothallus, such as found in many species <strong>of</strong> Fuscidea and Lecidella, is never formed. Small<br />
thalU forming ± circular patches amongst o<strong>the</strong>r lichens, suggesting parasitic behaviour, are<br />
characteristic <strong>of</strong> M. intrusa{p. 140).<br />
In vertical section <strong>the</strong> areolae usually lack a well-defined cortex, but <strong>the</strong>ir surface is sometimes<br />
covered by a thin (c. 2-10 (xm thick), hyaline amorphous layer which does not dissolve in<br />
concentrated KOH (Fig. lA, C). Such a layer is found in <strong>the</strong> areolae <strong>of</strong>, e.g. M. alabastrites, M.<br />
assimilata, M. cinerea, M. elachista, M. incrassata, M. lignaria, M. peliocarpa, M. subnigrata,<br />
M. subviolascens, and M. ternaria. Areolae with a similar external appearance, but without an<br />
amorphous covering layer, can be seen in well-developed specimens <strong>of</strong>, for example, M.<br />
denigrata, M. nitschkeana, M. globulosella, and sometimes A/, melaena (Fig. IB). The thallus <strong>of</strong><br />
<strong>the</strong>se species (and to a lesser extent that <strong>of</strong> M. elachista) is <strong>of</strong>ten invaded and disrupted by <strong>the</strong><br />
dematiaceous hyphae <strong>of</strong> a fungal parasite(s), and non-lichenized algae, resulting in a dark, <strong>of</strong>ten<br />
blackish, scurfy crust. This phenomen on is best exemplified by M. melaena, which in Britain is<br />
rarely found with a healthy, well-developed thallus. By contrast, during my excursions in<br />
Sweden I found it to be rarely parasitised. Susceptibility to invasion by parasites or opportunists<br />
may be related to climatic factors, being increased in <strong>the</strong> <strong>British</strong> Isles by <strong>the</strong> overall wea<strong>the</strong>r and<br />
warmer conditions. It would appear, <strong>the</strong>refore, that <strong>the</strong> amorphous covering layer (never<br />
present in M. melaena) is an effective barrier against potential invaders, but whe<strong>the</strong>r or not this<br />
is <strong>the</strong> reason for its evolution is a matter for speculation.<br />
The nearest approaches to <strong>the</strong> development <strong>of</strong> a true cortex are found in M. elachista and M.<br />
melaenida. Sections <strong>of</strong> healthy areolae show an outer layer (c. 10-12 /xm and 12-20 /xm thick<br />
respectively) <strong>of</strong> compacted, evacuated hyaline hyphae (Fig. IC). The amorphous covering layer<br />
<strong>of</strong> M. elachista <strong>of</strong>ten partially disintegrates causing <strong>the</strong> areolae to exhibit a white-pruinose<br />
appearance, a feature not noted in any o<strong>the</strong>r species <strong>of</strong> <strong>the</strong> genus.
LICHEN GENUS MICAREA IN EUROPE 23<br />
Fig. 1 Some thallus types in Micarea. A, areolate-type without cortex but with amorphous covering layer<br />
(e.g. LI lignaria). B, areolate-type without cortex or amorphous covering layer (e.g. M. denigrata). C,<br />
areolate-type with both cortex and amorphous covering layer (M. elachista). D, goniocysts (A/, prasina).<br />
Scale = 50 /Ltm.<br />
The outermost hyphae <strong>of</strong> areolae are frequently coloured or surrounded by a pigment , usually<br />
that which is found in <strong>the</strong> upper hymenium <strong>of</strong> <strong>the</strong> apo<strong>the</strong>cia, or <strong>the</strong> pycnidial walls, <strong>of</strong> <strong>the</strong> given<br />
species; for example, <strong>the</strong> dilute olivaceous, K+ violet pigment in M. denigrata, M. elachista, M.<br />
globulosella, M. nitschkeana, and M. subviolascens, <strong>the</strong> green, K— , HNO3+ red pigment in M.<br />
cinerea, M. lignaria, M. peliocarpa, M. sylvicola, and M. ternaria, and <strong>the</strong> brown, K- , HNO3—<br />
pigment in M. subnigrata. The intensity <strong>of</strong> pigmentation is much dependent on exposure to light<br />
and pigment may be entirely absent in shade forms. Species that always lack pigment in <strong>the</strong>ir<br />
apo<strong>the</strong>cia and pycnidia (e.g. M. alabastrites, M. pycnidiophora, amd M. stipitata) similarly lack<br />
pigment in <strong>the</strong> thallus.<br />
In <strong>the</strong> areolae <strong>of</strong> most species <strong>the</strong> algal layer is in direct contact with <strong>the</strong> substratum. However,<br />
a white medulla, devoid <strong>of</strong> algal cells, may be formed in <strong>the</strong> larger areolae <strong>of</strong> some species, e.g.<br />
M. incrassata, M. intrusa, M. lignaria, and M. subnigrata.
24 BRIAN JOHN COPPINS<br />
(ii) Goniocyst-type (Fig. ID)<br />
Several species have a finely granular thallus composed <strong>of</strong> discrete, ± globular structures,<br />
mostly c. 12-40 /am diam. These ecorticate granules consist <strong>of</strong> clustered algal cells intertwined<br />
and surrounded by short-celled hyphae, and are never protected by an amorphous covering<br />
layer. They are <strong>of</strong>ten seen to have short protruding hyphae, but <strong>the</strong>y never have distinct spines<br />
as found in some species <strong>of</strong> Vezdaea (Poelt & Dobbeler, 1975). They are similar to soredia, but<br />
because <strong>the</strong>y are <strong>the</strong> main component <strong>of</strong> <strong>the</strong> thallus and not derived from specialised structures<br />
(soralia) or eroding or disintegrating parts <strong>of</strong> a different thallus type (see (iii) below), <strong>the</strong> term<br />
goniocyst (Ozenda, 1963) seems <strong>the</strong> most applicable for <strong>the</strong>m.<br />
In M. prasina, M. hedlundii, and M. melanobola <strong>the</strong> thallus is composed entirely <strong>of</strong><br />
goniocysts, <strong>the</strong> first <strong>of</strong> which appear to arise directly from <strong>the</strong> prothalline hyphae; fur<strong>the</strong>r<br />
development is presumably by a process <strong>of</strong> division or budding from existing goniocysts. M.<br />
botryoides has a 'primary' thallus <strong>of</strong> flattened granular-areolae and in a few specimens I have<br />
noticed goniocysts developing from <strong>the</strong>se areolae, which <strong>the</strong>n become obscured as <strong>the</strong> gonio-<br />
cysts proliferate. However, in most specimens <strong>of</strong> M. botryoides no areolae can be seen; ei<strong>the</strong>r<br />
<strong>the</strong>y have become obscured or, perhaps in some cases <strong>the</strong> thallus develops as goniocysts from <strong>the</strong><br />
outset.<br />
In M. prasina and M. melanobola <strong>the</strong> outermost hyphae <strong>of</strong> superficial (exposed) goniocysts<br />
are <strong>of</strong>ten surrounded by <strong>the</strong> K+ violet pigment which also occurs in <strong>the</strong> upper hymenium <strong>of</strong><br />
<strong>the</strong>se species. When well developed, thalli composed <strong>of</strong> goniocysts have a ± gelatinous<br />
appearance when moist; this was <strong>the</strong> reason why Elias Fries originally placed <strong>the</strong> genus Micarea<br />
in his 'Tribus Collemaceae'.<br />
The thallus <strong>of</strong> M. syno<strong>the</strong>oides consists <strong>of</strong> small dark granular-areolae and is somewhat<br />
intermediate between <strong>the</strong> 'areolate-type'. The thallus <strong>of</strong> M. myriocarpa sometimes has a scurfy<br />
appearance but is <strong>of</strong>ten organized into small (10-15 /am diam) goniocyst-like granules.<br />
(iii) Sorediate-type<br />
This is represented by <strong>the</strong> apparently closely related M. leprosula and M. subleprosula. Their<br />
thalli are essentially <strong>of</strong> <strong>the</strong> 'areolate-type' but <strong>the</strong>ir areolae lack an amorphous covering layer<br />
and are very fragile (easily broken by touching with <strong>the</strong> point <strong>of</strong> a needle), <strong>of</strong>ten breaking<br />
down or eroding to form irregularly shaped, pale green or yellowish green soredial granules,<br />
c. 20-50 /Ltm diam.<br />
(iv) Smooth- or scurfy-type<br />
Included here are thalli which, although developed on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> substratum, are not<br />
organized into discrete areolae or goniocysts. The thalh may be smooth, continuous to rimose,<br />
or irregularly scurfy-granular. Such thalli may be formed by species that normally have well<br />
defined areolae (e.g. M. lignaria and M. melaenida) or, by lignicolous species whose thalli are<br />
<strong>of</strong>ten endoxylic (e.g. M. anterior, M. nigella, and M. olivacea). The thalli <strong>of</strong> M. bauschiana and<br />
M. lutulata vary from smooth or rimose, to scurfy-granular, and lack areolae, whereas <strong>the</strong><br />
similar M. lithinella, M. sylvicola, and M. tuberculata sometimes produce areolae. Ano<strong>the</strong>r two<br />
species whose thalli fall within this category are M. adnata and M. turfosa.<br />
(v) Immersed-type<br />
The thallus <strong>of</strong> many lignicolous species is indistinct and developed below <strong>the</strong> surface <strong>of</strong> <strong>the</strong><br />
substratum, which may have a bleached appearance (cf. M. muhrii). An endoxylic thallus is<br />
occasionally formed by species that more characteristically have superficial areolae, e.g. M.<br />
cinerea, M. denigrata, M. lignaria, M. muhrii, and M. peliocarpa. Conversely, M. melaeniza and<br />
M. misella are usually endoxyUc but forms with areolae are sometimes encountered. The thallus<br />
in M. anterior, M. nigella, and M. olivacea is <strong>of</strong>ten endoxylic, but a thin ± smooth crust is<br />
sometimes formed on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> lignum.<br />
The thalli <strong>of</strong> M. contexta, M. eximia, and M. rhabdogena are invariably endoxylic. Sections <strong>of</strong><br />
<strong>the</strong> substratum show small goniocyst-like clusters (c. 15-40 /am diam) surrounded and interconnected<br />
by pigmented hyphae. As in <strong>the</strong> relevant examples in thallus types (i-iii), and in M.<br />
turfosa in group (iv), <strong>the</strong> pigment involved is that found in <strong>the</strong> upper hymenium or pycnidial<br />
walls <strong>of</strong> <strong>the</strong> given species, viz. : dark green and K— in M. contexta and M. eximia, and olivaceous<br />
and K-f- violet in M. rhabdogena.
LICHEN GENUS MICAREA IN EUROPE 25<br />
The species which at times grow on bryophytes usually have a superficial thallus, but even so,<br />
<strong>the</strong> thallus is <strong>of</strong>ten, in part, endocuticular. When on bark Micarea thalli are generally<br />
epiphloeodal, but at least partially endophloeodal thalli have been seen in M. cinerea and M.<br />
peliocarpa.<br />
Phycobionts<br />
Among <strong>the</strong> 45 species <strong>of</strong> Micarea accepted in this revision three types (? genera) <strong>of</strong> 'grass-green'<br />
phycobiont can be distinguished by LM observations on thallus squashes. Unfortunately <strong>the</strong><br />
identities <strong>of</strong> <strong>the</strong>se algae are unknown and <strong>the</strong>y await critical studies by an algologist. In addition,<br />
two genera <strong>of</strong> blue-green alga may be involved in <strong>the</strong> formation <strong>of</strong> cephalodia (^.v.).<br />
The commonest alga in Micarea is that found in <strong>the</strong> type species {M. prasina) and was<br />
discussed at some length by Hedlund (1892, 1895). In <strong>the</strong> course <strong>of</strong> this study this alga is<br />
hereafter referred to as 'micareoid' (Fig. 2). Its cells are fairly regular in appearance, ± globose,<br />
thin-walled and c. 4-7 jxm diam. Reproduction within <strong>the</strong> thallus appears to be by a process <strong>of</strong><br />
cell division in which <strong>the</strong> protoplast divides in two, followed by <strong>the</strong> laying down <strong>of</strong> a dividing wall<br />
which joins up with existing wall <strong>of</strong> <strong>the</strong> parent cell (Fig. 2A). No internal divisions into<br />
aplanospores as found in, for example, Myrmecia and Trebouxia, have been observed. Contact<br />
between <strong>the</strong> fungal hyphae and <strong>the</strong> algae cells is by intracellular haustoria (Peveling, 1974)<br />
which are readily seen at x 1000 (mounted in 10% KOH, followed by ammoniacal erythrosin).<br />
The haustoria may be peg-like (Fig. 2B, D-E), swollen to become clavate (Fig. 2G) or capitate<br />
(Fig. 2C, H), or spreading into a foot (Fig. 2F). In collapsed algal cells (Fig. 2H) <strong>the</strong> haustoria<br />
are intensively stained. A characteristic feature <strong>of</strong> <strong>the</strong> micareoid alga and its relationship with<br />
<strong>the</strong> mycobiont is that a hypha frequently becomes closely aligned along <strong>the</strong> dividing line <strong>of</strong> two<br />
separating algal daughter cells (Fig. 2D-E); <strong>the</strong> hypha concerned is <strong>of</strong>ten seen to penetrate one<br />
or both <strong>of</strong> <strong>the</strong> cells by haustoria.<br />
The second algal type is found in Micarea sylvicola and its presumed relatives M. bauschiana,<br />
M. lutulata, and M. tuberculata. The cells are thin-walled and irregular in size and shape, varying<br />
from 5-12 /xm diam when globose to up to c. 15 x 10 /xm when ± ellipsoid. I am undecided as to<br />
<strong>the</strong>ir mode <strong>of</strong> reproduction within <strong>the</strong> thallus. The large number <strong>of</strong> cells at <strong>the</strong> lower end <strong>of</strong> <strong>the</strong><br />
size range is suggestive <strong>of</strong> aplanospore formation, but I have never observed a mo<strong>the</strong>r-cell<br />
undergoing division. Haustorial penetrations by <strong>the</strong> mycobiont hyphae have been detected, but<br />
<strong>the</strong>y are never as distinct as those involving micareoid algae or <strong>the</strong> phycobiont <strong>of</strong> M. intrusa (see<br />
below).<br />
The third algal type is confined to M. intrusa, and has large globose cells, 7-21 /^m diam, with<br />
thick hyaline walls c. l-lfxm thick. Cell division within <strong>the</strong> thallus seems to be by a process <strong>of</strong> <strong>the</strong><br />
division <strong>of</strong> a mo<strong>the</strong>r-cell into three or four daughter-cells (i.e. 'protococcoid' division). Many <strong>of</strong><br />
<strong>the</strong> cells are clearly seen to be deeply penetrated by a haustorium (Fig. 55). This phycobiont<br />
looks very similar to that <strong>of</strong> Scoliciosporum umbrinum and may well be identical with it.<br />
Cephalodia<br />
For reviews <strong>of</strong> <strong>the</strong> morphological, taxonomic, and physiological aspects <strong>of</strong> cephalodia see Jahns<br />
(1974), James & Henssen (1976), and Millbank (1976).<br />
Cephalodia have been found in three species <strong>of</strong> Micarea: M. assimilata, M. incrassata and M.<br />
subviolascens . The first two species have thalli composed <strong>of</strong> verrucose areolae with a micareoid<br />
phycobiont. However, sections <strong>of</strong> <strong>the</strong>ir thalli reveal <strong>the</strong> presence <strong>of</strong> ± globose structures (c.<br />
200-600 /xm diam) containing a blue-green alga <strong>of</strong> <strong>the</strong> genus Nostoc. In many cases <strong>the</strong>se<br />
structures are visible externally and closely resemble <strong>the</strong> areolae except that <strong>the</strong>y are brown, and<br />
usually darker, in colour. Internally <strong>the</strong>y consist <strong>of</strong> numerous ramifying fungal hyphae (presumably<br />
belonging to <strong>the</strong> Micarea) and dense masses <strong>of</strong> Nostoc cells which have lost <strong>the</strong>ir normal<br />
(when free-living) filamentous arrangements; and I am in no doubt that <strong>the</strong>se structures are<br />
cephalodia. I am less certain <strong>of</strong> <strong>the</strong> status <strong>of</strong> <strong>the</strong> more loosely organized clusters <strong>of</strong> Stigonema<br />
which are sometimes associated with <strong>the</strong> same two Micarea species, and also M. subviolascens.<br />
However, <strong>the</strong> Stigonema filaments are, at least partially, disrupted and fungal hyphae are<br />
present.
26 BRIAN JOHN COPPINS<br />
A B<br />
Fig. 2 'Micareoid' phycobiont <strong>of</strong> Micarea alabastrites and its relationship with mycobiont hyphae ; see text<br />
for fur<strong>the</strong>r details. Scale = 10 /xm.<br />
The three above-mentioned Micarea species are morphologically closely related and occur in<br />
exposed arctic-alpine situations. In such habitats <strong>the</strong> formation <strong>of</strong> cephalodia with a blue-green<br />
alga(e) which has <strong>the</strong> ability to fix atmospheric nitrogen, is <strong>of</strong> undoubted nutritional benefit to<br />
<strong>the</strong> mycobionts. Most morphological and physiological studies concerning cephalodia have<br />
involved macrolichens. Reports <strong>of</strong> cephalodia in crustose lichen are relatively few, although<br />
some are Hsted by James & Henssen (1976). Additional examples in <strong>the</strong> Lecideaceae include<br />
Huilia aeolotera, H. elegantior, H. panaeola (Hertel, 1977), and Lecidea pallida (Fries, 1874).<br />
Apo<strong>the</strong>cia: external features<br />
As a rule <strong>the</strong> apo<strong>the</strong>cia in Micarea are small to medium sized, convex to ± globose or<br />
tuberculate, and usually immarginate. However, this generalization encompasses considerable<br />
variation even within a single given species. With regard to shape, some species (e.g. M.<br />
denigrata and M. prasina) are very variable with apo<strong>the</strong>cia that may be shallow-convex to<br />
convex-hemispherical (Figs 3A-B, 4A-B) and finally ± globose (Fig. 3D) or tuberculate<br />
(Fig. 3C). Apo<strong>the</strong>cia <strong>of</strong> o<strong>the</strong>r species may be less variable: those <strong>of</strong> M. alabastrites and M.<br />
cinerea are broadly convex to hemispherical, sometimes tuberculate, but never ± globose; at <strong>the</strong><br />
H
LICHEN GENUS MICAREA IN EUROPE 27<br />
^>t.<br />
OQ<br />
LU<br />
3 3
28 BRIAN JOHN COPPINS<br />
3 00
LICHEN GENUS MICAREA IN EUROPE 29<br />
O<strong>the</strong>r extreme those <strong>of</strong> M. contexta, M. eximia, and M. myriocarpa are ± globose from <strong>the</strong><br />
beginning and <strong>of</strong>ten tuberculate . The range <strong>of</strong> variabihty for most o<strong>the</strong>r species falls somewhere<br />
between <strong>the</strong>se last two extremes. Apo<strong>the</strong>cia are usually sessile (or partly hidden by adjoining<br />
areolae or goniocysts) but occasionally, as with M. lignaria, M. botryaides, and M. ternaria, <strong>the</strong>y<br />
may be turbinate or short-stalked (due to vertical elongation <strong>of</strong> excipular and (or) hypo<strong>the</strong>cial<br />
tissues), but only in M. crassipes and an undescribed species from New Zealand is this a constant<br />
feature (Figs 3E, 4B). Apo<strong>the</strong>cia are usually immarginate, but in those species with a well<br />
differentiated excipulum (e.g. M. cinerea, M. denigrata, M. muhrii, M. peliocarpa, and M.<br />
ternaria), a faint marginal zone is sometimes apparent, especially in young apo<strong>the</strong>cia. This zone<br />
is rarely raised above <strong>the</strong> level <strong>of</strong> <strong>the</strong> disc, but it is <strong>of</strong>ten paler in colour (especially when wet) and<br />
slightly more glossy. Even in M. crassipes, <strong>the</strong> only species whose apo<strong>the</strong>cia regularly exhibit a<br />
distinctly raised marginal rim, this feature is soon occluded as <strong>the</strong> disc expands and becomes<br />
convex.<br />
The dimensions <strong>of</strong> apo<strong>the</strong>cia in Micarea fall mainly within <strong>the</strong> range 0- 1-0-6 mm diam. In<br />
some species (e.g. M. denigrata, M. peliocarpa, and M. prasina) <strong>the</strong>re is considerable variation<br />
in apo<strong>the</strong>cial size, but in o<strong>the</strong>rs it may be less so. Characteristically small apo<strong>the</strong>cia (
30 BRIAN JOHN COPPINS<br />
Fig. 5 Sketch demonstrating a situation which M. bauschiana shows variation in <strong>the</strong> colour <strong>of</strong> its<br />
apo<strong>the</strong>cia, from paUid (A), through pallid/brownish or pallid/blue-grey (B), to blue-black (C). Site:<br />
roadside bank in woodland in lower Glen Roy, Westerness (cf. Coppins 3492/3, E); <strong>the</strong> bank is c. 1 m<br />
high).<br />
M. denigrata, M. nitschkeana, and M. prasina. I know <strong>of</strong> no Micarea that has apo<strong>the</strong>cia<br />
possessing pruina or, exhibiting a bluish (caesious) 'bloom' when wetted. Such features are due<br />
to <strong>the</strong> existence in <strong>the</strong> epi<strong>the</strong>cium <strong>of</strong> minute colourless (or pale straw when dense) granules that<br />
dissolve in K, and are characteristic <strong>of</strong> several species <strong>of</strong>ten misidentified to Micarea, e.g.<br />
Bacidia beckhausii, Lecanora symmicta agg. , Lecidea caesioatra, L. turgidula, and Scoliciosporumpruinosum.<br />
Apo<strong>the</strong>cia: internal features<br />
All tissues <strong>of</strong> <strong>the</strong> apo<strong>the</strong>cia are bound by a weak gel-matrix which ± dissipiates in 10% KOH to<br />
clearly reveal <strong>the</strong> component parts (i.e. hyphae, paraphyses, and asci). In some species <strong>the</strong><br />
apices <strong>of</strong> <strong>the</strong> paraphyses and/or <strong>the</strong> hyphae <strong>of</strong> <strong>the</strong> hypo<strong>the</strong>cium are additionally cemented by<br />
dense pigment deposited in <strong>the</strong> matrix and/or on <strong>the</strong> hyphal walls, thus tending to impair <strong>the</strong>ir<br />
separation in K. The presence <strong>of</strong> <strong>the</strong> gel-matrix serves to separate species <strong>of</strong> Micarea from some<br />
superficially similar species <strong>of</strong> Vezdaea (Poelt & Dobbeler, 1975).<br />
Hymenium and epi<strong>the</strong>cium<br />
The term 'epi<strong>the</strong>cium' is here employed in its broad sense to refer to <strong>the</strong> upper part (usually c.<br />
3-15 /xm) <strong>of</strong> <strong>the</strong> hymenium where this differs in appearance from <strong>the</strong> remaining lower part(s).
LICHEN GENUS MICAREA IN EUROPE 31<br />
This difference can be due to <strong>the</strong> much branched and entangled apices <strong>of</strong> <strong>the</strong> paraphyses, or<br />
(and) deposits <strong>of</strong> pigment in <strong>the</strong> gel-matrix, or on (or in) <strong>the</strong> walls <strong>of</strong> <strong>the</strong> paraphyses. The upper<br />
layer <strong>of</strong> <strong>the</strong> hymenium differentiated in such ways is perhaps more correctly termed <strong>the</strong><br />
'epihymenium' (Poelt, 1974a) or 'pseudoepi<strong>the</strong>cium' (Korf, 1973). In its strict (original) sense<br />
an 'epi<strong>the</strong>cium' refers to a layer <strong>of</strong> branches <strong>of</strong> paraphyses that overtop <strong>the</strong> asci. Such a layer is<br />
approached in some species <strong>of</strong> Micarea and is best exemplified by M. contexta and M.<br />
melanobola. In Micarea <strong>the</strong> difference between an 'epihymenium' and an 'epi<strong>the</strong>cium' is not<br />
clear-cut, and so I have chosen to employ <strong>the</strong> latter term which has been long and widely<br />
employed in lichenology.<br />
The height <strong>of</strong> <strong>the</strong> hymenium in Micarea species is sometimes difficult to measure accurately<br />
because <strong>the</strong> hymenium <strong>of</strong>ten merges ra<strong>the</strong>r imperceptibly into <strong>the</strong> hypo<strong>the</strong>cium. Measurements<br />
are most accurately made by mounting thin sections in Lugol's iodine, in which <strong>the</strong> strongly<br />
amyloid (dark blue) hymenium contrasts strongly with <strong>the</strong> non-amyloid (or ± so) hypo<strong>the</strong>cium.<br />
Even when accurately determined <strong>the</strong> height <strong>of</strong> <strong>the</strong> hymenium is rarely a useful character in<br />
distinguishing similar species. The overall range in height is about 23-90 /am, and in most cases is<br />
within 35-50 /xm. Shallow hymenia (rarely exceeding 35 /am) are characteristic <strong>of</strong> many <strong>of</strong> <strong>the</strong><br />
species with small spores, viz.: M. hedlundii, M. melanobola, M. misella, M. myriocarpa, M.<br />
nigella, M. olivacea, M. osloensis, M. rhabdogena, and M. tuberculata. Likewise, tall hymenia<br />
are characteristic <strong>of</strong> species with large spores, and <strong>the</strong> tallest hymenium (65-90 /u,m) belongs to<br />
<strong>the</strong> species with <strong>the</strong> largest spores, namely M. subleprosula. The height <strong>of</strong> <strong>the</strong> hymenium<br />
sometimes increases with <strong>the</strong> age <strong>of</strong> <strong>the</strong> apo<strong>the</strong>cium, and this is particularly true in M.<br />
bauschiana and M. sylvicola: small, yet mature, apo<strong>the</strong>cia usually have a hymenium c. 40 /zm<br />
tall, but as <strong>the</strong> apo<strong>the</strong>cium enlarges and increases in convexity, <strong>the</strong> hymenium <strong>of</strong>ten increases in<br />
height up to about 60 /xm; at <strong>the</strong> same time <strong>the</strong> paraphyses appear to 'stretch' and become<br />
thinner (especially in <strong>the</strong> lower part <strong>of</strong> <strong>the</strong> hymenium).<br />
The hymenium in Micarea never contains hyaline, crystalline inclusions that adhere to<br />
paraphyses (as found in some species <strong>of</strong> Lecidella, and many Graphidaceae) or numerous oil<br />
droplets (as found in several species <strong>of</strong> Buellia and Caloplaca). Epi<strong>the</strong>cial granules that dissolve<br />
in K and <strong>of</strong>ten give <strong>the</strong> apo<strong>the</strong>cia a pruinose appearance are also never found in Micarea. Minute<br />
granules <strong>of</strong> blue-violet (K+aeruginose) pigment, like those found in Lecidea hypnorum, are<br />
occasionally observed in <strong>the</strong> hymenium <strong>of</strong> M. contexta and M. lignaria.<br />
The colour <strong>of</strong> <strong>the</strong> hymenium and epi<strong>the</strong>cium is an important character in <strong>the</strong> identification <strong>of</strong><br />
Micarea species. A discussion <strong>of</strong> <strong>the</strong> various pigments involved is given in <strong>the</strong> sections on<br />
'chemistry'. The hymenium may be dilutely and ± evenly coloured throughout, or <strong>the</strong><br />
colouration may be more intense in <strong>the</strong> upper part. In many species <strong>the</strong> hymenium is <strong>of</strong>ten<br />
intersected by dark vertical streaks, a feature usually due to dense pigment surrounding<br />
individual, or small fascicles <strong>of</strong> stout paraphyses (see p. 61).<br />
Asci<br />
The asci <strong>of</strong> Micarea species are clavate or cylindrical-clavate in shape and belong to <strong>the</strong><br />
Lecanora-type <strong>of</strong> Honegger (1978). When mounted in Lugol's iodine (following treatment in<br />
10% KOH) <strong>the</strong> ascus is seen to have a non-amyloid wall surrounded by an amyloid outer-layer<br />
('fuzzy coat') and an internal staining amyloid apical dome (Fig. 6). Micarea has sometimes<br />
been placed in <strong>the</strong> Arthoniaceae (e.g. Lamb, 1953) but <strong>the</strong> ascus <strong>of</strong> members <strong>of</strong> that family does<br />
not have an amyloid outer-layer and <strong>the</strong> apical dome is not deeply amyloid. In a few species<br />
<strong>of</strong> Arthonia and Artho<strong>the</strong>lium I have observed a faint bluing in <strong>the</strong> part <strong>of</strong> <strong>the</strong> apical dome<br />
immediately adjacent to <strong>the</strong> ocular chamber. In some Arthonia species a tiny amyloid ring is<br />
apparent above <strong>the</strong> apex <strong>of</strong> <strong>the</strong> ocular chamber, and, despite statements to <strong>the</strong> contrary by<br />
Eriksson (1981), such a ring is found in <strong>the</strong> asci <strong>of</strong> ^. radiata (type species oi Arthonia Ach.) and<br />
A. fuscopurpurea. The same type <strong>of</strong> ring was also described for Bryostigma leucodontis by Poelt<br />
& Dobbeler (1979). Such aspects <strong>of</strong> ascus structure in <strong>the</strong> Arthoniaceae clearly merit more<br />
detailed investigation. The ascus structure in Micarea supports my opinion that <strong>the</strong> genus should<br />
be placed in <strong>the</strong> Lecideaceaes. str., at least for <strong>the</strong> time being.<br />
The asci <strong>of</strong> all European Micarea species are 8-spored. The North American Lecidea populina
32 BRIAN JOHN COPPINS<br />
Fig. 6 Ascus apex <strong>of</strong> Micarea alabastrites in optical section (LM); mounted in Lugol's iodine following<br />
pre-treatment in 10% KOH. A, young ascus. B, mature ascus. ac, apical cushion; aw, ascus wall; d,<br />
apical dome (tholus); oc, ocular chamber; ol, outer layer <strong>of</strong> ascus wall. Shading indicates intensity <strong>of</strong><br />
amyloid reaction; note that in reality <strong>the</strong> apical cushion and ocular chamber are probably completely<br />
non-amyloid. Scale = 10 /xm.<br />
Miill. Arg. has 16-spored asci and was transferred to Micarea by Anderson & Carmer (1974).<br />
Unfortunately <strong>the</strong> type material <strong>of</strong> L. populina has been out on loan from H and not available to<br />
me during <strong>the</strong> course <strong>of</strong> this study. However, <strong>the</strong> species apparently occurs in Xanthorion<br />
communities and, if this is so, <strong>the</strong>n it is unUkely to be a Micarea.<br />
Spores<br />
The wide variety <strong>of</strong> spore types found in Micarea can be seen in <strong>the</strong> selection <strong>of</strong> spores from all<br />
<strong>the</strong> European species illustrated in Figs 7-33. A few species (e.g. M. assimilata, M. contexta, and<br />
M. lithinella) have spores that are ± consistent in size, shape, and septation, but for many o<strong>the</strong>r<br />
species (e.g. M. anterior, M. botryoides, M. denigrata, M. prasina, and M. turfosa) <strong>the</strong>se<br />
characters can be very variable even within <strong>the</strong> same ascus. The smallest spores are found in M.<br />
myriocarpa (5-5-8-5xl-5-2-5 jxm) and <strong>the</strong> largest ones are found in M. subleprosula (40-60X<br />
5-6-6 /Ltm). Spores may be simple or up to 7-septate, <strong>the</strong> variations within <strong>the</strong>se hmits depending<br />
on <strong>the</strong> species. Spores with more than 7-septa are very rare, although a few 9-septate spores have<br />
been observed in M. subleprosula, and a collection (Coppins, 1834) <strong>of</strong> M. syno<strong>the</strong>oides has<br />
spores with up to 11 septa. Among <strong>the</strong> great range <strong>of</strong> spore shapes encountered, some may be<br />
broadly ellipsoid {M. subnigrata and M. intrusa) ,<br />
but regularly globose spores do not occur in<br />
Micarea.<br />
Healthy spores are always thin-walled and colourless. However, <strong>the</strong> walls <strong>of</strong> old spores<br />
trapped within <strong>the</strong> hymenium sometimes become slightly thickened and pale straw coloured, or<br />
<strong>the</strong>y may become impregnated with hymenial pigment. Spore walls always appear smooth (LM<br />
at X 1000) and are never surrounded by a gelatinous epispore. The cytoplasm <strong>of</strong> spores, asci, and<br />
ascogenous hyphae in M. intrusa is sometimes a dilute orange, turning purple-red in K.
LICHEN GENUS MICAREA IN EUROPE 33<br />
Fig. 7 A, M. adnata (E - holotype). B, M. anterior (H-NYL 21655 - lectotype). Scale = 10 /xm.<br />
B
34 BRIAN JOHN COPPINS<br />
Fig. 8 M. alabastrites. A, (H-NYL 18656 - holotype). B, {Coppins 2588, E). Scale = 10 /ixm.<br />
B
h<br />
LICHEN GENUS MJCAREA IN EUROPE 35<br />
Fig. 9 M. assimilata. A, (H-NYL 16556 -lectotype). B, (Norway, Oppland, Fogstuen, 1857, Lindsay, E).<br />
Scale = 10 /xm.<br />
B
36 BRIAN JOHN COPPINS<br />
Fig. 10 A, M. bauschiana (M - lectotype). B, M. botryoides {Coppins 8429, E). Scale = 10 /xm.
LICHEN GENUS MICAREA IN EUROPE 37<br />
I- H<br />
Fig. 11 M. cinerea {Coppins 2533, E). Scale = 10 /Ltm.
38 BRIAN JOHN COPPINS<br />
Fig. 12 A,M. contexta (S-\ectotype). B,M. crassipes (VezdaLich. Sel. 11,BM).C,A/. cMrvara (WRSL<br />
holotype). Scale = 10 fxm.<br />
H<br />
B<br />
-
LICHEN GENUS MICAREA IN EUROPE 39<br />
f]r\<br />
VJ u<br />
Fig. 13 M. denigrata. A, (UPS - lectotype). B, (H - lectotype <strong>of</strong> Lecidea hemipoliella). Scale = 10 ^tm.<br />
B
40 BRIAN JOHN COPPINS<br />
r\<br />
U<br />
Fig. 14 A, M. elachista (L - lectotype). B, M. eximia (S - lectotype). C, M. hedlundii (UPS - holotype).<br />
Scale = 10 /xm.
LICHEN GENUS MICAREA IN EUROPE 41<br />
Fig. 15 M. globulosella. A, (S - lectotype). B, (Czechoslovakia, Slovakia, Vysoke Tatry, 1879, Lojka,<br />
BM). Scaler 10 /am.<br />
y
42 BRIAN JOHN COPPINS<br />
Fig. 16 M. incrassata. A, (S - holotype). B, (Kerguelen, 1875, Eaton, E). Scale = 10 /u,m.<br />
B
LICHEN GENUS MICAREA IN EUROPE 43<br />
Fig. 17 M. intrusa. A, (UPS - holotype). B, (BM - isotype <strong>of</strong> Lecidea melaphana). C, (Norway,<br />
Hordaland, Fjell, 1980, Skjolddal, BG). Scale = 10 /xm.
44 BRIAN JOHN COPPINS<br />
Fig. 18 M. leprosula (UPS-lectotype). Scale — 10)u,m.
LICHEN GENUS MICAREA IN EUROPE 45<br />
Fig. 19 M. lignaria. A, (H-ACH 265 - lectotype). B, (TUR-VAINIO 21274- lectotype <strong>of</strong> Lecidea trisepta<br />
yar.polytropoides). Scale = 10 ixm.<br />
H
46 BRIAN JOHN COPPINS<br />
Fig. 20 A-B, M. lithinella; A, (M - lectotype) ; B, (Germany, Heidelberg, Zwdckh, H-NYL 19191). C, M.<br />
lutulata (Wales, Pembroke, Tycanol, 1980, James, BM). Scale = 10 ^tm.<br />
B
LICHEN GENUS MICAREA IN EUROPE 47<br />
B<br />
Fig. 21 A, M. melaena (UPS - lectotype <strong>of</strong> Lecidea milliaria var. turfosa). B, M. melaeniza (S - holotype)<br />
C, M. melanobola (H-NYL 21614 - lectotype). Scale = 10 /Ltm.<br />
.
48<br />
BRIAN JOHN COPPINS<br />
Fig. 22 M. melaenida. A, (H-NYL 18843 - holotype). B, (Fl. Hung. exs. 714, BM - topotype <strong>of</strong> Catillaria<br />
zsakii). C, {^'RSL-\ec\o\.yptoi Catillaria schumanii). Scale = lOjLtm.<br />
B
LICHEN GENUS MICAREA IN EUROPE 49<br />
Fig. 23 A, M. misella (H-ACH 52-holotype <strong>of</strong> Lecidea resinae subsp. globularis). B, M. muhrii (E<br />
holotype). C, M. myriocarpa (hb Wirth 6085 - holotype). Scale = 10 /u,m.<br />
B
50 BRIAN JOHN COPPINS<br />
Fig. 24 A, M. nigella (E - holotype). B, M. nitschkeana (Coppins 2426, E). C, M. olivacea (BM<br />
holotype). Scale = 10/u,m.<br />
B
LICHEN GENUS MICAREA IN EUROPE 51<br />
Fig. 25 M. peliocarpa. A, (H-NYL 18716 - isotype <strong>of</strong> Lecidea violacea). B, (England, New Forest, Great<br />
Wood, Bramble Hill Walk, 1970, Coppins et al. , E). Scale = 10 />tm.<br />
B
52 BRIAN JOHN COPPINS<br />
Fig. 26 M. prasina. A, (UPS - lectotype). B, (H-NYL 19056 - lectotype <strong>of</strong> Lecidea subviridescens). Scale<br />
= 10 /um.<br />
B
LICHEN GENUS MICAREA IN EUROPE 53<br />
Fig. 27 M. prasina. A, (H-NYL 21604 - lectotype <strong>of</strong> Lecidea prasiniza) . B, (GZU - holotype <strong>of</strong> Micarea<br />
polytrichi). Scale = 10 /xm.<br />
B
54 BRIAN JOHN COPPINS<br />
VJ<br />
Fig. 28 A, M. pycnidiophora (E - holotype). B, M. osloensis (UPS - holotype). Scale = 10 ju-m.<br />
B
LICHEN GENUS MICAREA IN EUROPE 55<br />
r\<br />
Fig, 29 A,M. rhabdogena (BM - isolectotype). B, M. stipitata (E - holotype). Scale = 10 /u-m.<br />
\j
56 BRIAN JOHN COPPINS<br />
Fig. 30 M. subleprosula (Mu/ir 4380, E). Scale = 10 /Am.
LICHEN GENUS MICAREA IN EUROPE 57<br />
Fig. 31 A, M. subnigrata (H-NYL 19136 -lectotype). B, M. subviolascens (Havaas Lich. Exs. Norv. 139,<br />
BG). C, M. sylvicola (UPS - lectotype). D-E, M. syno<strong>the</strong>oides; D, (H-NYL 19101 - lectotype); E,<br />
(Coppins 3257, E). Scale = 10 ixm.<br />
Fig . 3 1 D-E on foliowing page<br />
B
58<br />
BRIAN JOHN COPPINS<br />
Fig. 31 - cont.
LICHEN GENUS MICAREA IN EUROPE 59<br />
^<br />
Fig. 32 A, M. tuberculata (O - lectotype). B, M. ternaria {Thomson 9188, DUKE). Scale = 10 /xm.<br />
r\<br />
u
60 BRIAN JOHN COPPINS<br />
Fig. 33 M. turfosa. A, (VER- holotype). B, (Vezda Lich. Sel. 1135, BM). Scale = 10 /xm.
LICHEN GENUS MICAREA IN EUROPE 61<br />
Paraphyses<br />
The paraphyses in Micarea are characteristically thin and branched. When measured (in 10%<br />
KOH, or ammoniacal erythrosin) at <strong>the</strong> mid-hymenium <strong>the</strong>y may be very thin and only about<br />
0-6-1 fxm wide (e.g. M. anterior, M. botryoides, M. contexta, M. eximia, M. lithinella, M.<br />
misella, and M. prasina), or thin and c. 1-1-5 /xm wide (e.g. M. adnata, M. cinerea, M. denigrata,<br />
M. intrusa, M. muhrii, and M. peliocarpa), or relatively stout and about 1-5-1-8 fjun wide (e.g.<br />
M. assimilata, M. incrassata, M. lignaria, M. osloensis, and M. subnigrata) ; <strong>the</strong>se measurements<br />
relate to paraphyses not coated in pigment. In old, much expanded, apo<strong>the</strong>cia <strong>the</strong> paraphyses<br />
sometimes appear 'stretched' and thinner than normal (especially in <strong>the</strong> lower half <strong>of</strong> <strong>the</strong><br />
hymenium) ; this phenomenon has frequently been observed in collections <strong>of</strong> M. bauschiana, M.<br />
sylvicola, and M. lignaria. In many species <strong>the</strong> paraphyses gradually widen towards <strong>the</strong>ir apices,<br />
but <strong>the</strong> apices are never regularly clavate or capitate. This widening is <strong>of</strong>ten enhanced by <strong>the</strong><br />
deposition <strong>of</strong> closely adhering pigment which sometimes gives <strong>the</strong> appearance <strong>of</strong> a 'hood' (e.g.<br />
M. melaena) ; such coatings or hoods can be detached by gently boiling and <strong>the</strong>n tapping sections<br />
or squash preparations in 50% KOH. In a few cases (e.g. M. melanobola) <strong>the</strong> pigment cannot be<br />
separated in this way and appears to be located within <strong>the</strong> walls <strong>of</strong> <strong>the</strong> paraphyses. Paraphyses<br />
with dark pigmented apical 'caps', like those found in Catillaria s. str. (Killias, 1980: 253),<br />
Buellia, and many species <strong>of</strong> Lecanora, are not known in Micarea.<br />
In all species <strong>of</strong> Micarea a large proportion <strong>of</strong> <strong>the</strong> paraphyses are branched, even if <strong>the</strong><br />
branching is mainly confined to <strong>the</strong> epi<strong>the</strong>cium. Species with sparingly branched paraphyses<br />
include M. assimilata, M. incrassata and M. lignaria. Anastomozing paraphyses have been<br />
observed in all <strong>the</strong> species, but <strong>of</strong>ten <strong>the</strong> anastomoses are ± confined to <strong>the</strong> lower third <strong>of</strong> <strong>the</strong><br />
hymenium. The degree <strong>of</strong> branching and anastomosing is difficult to quantify for <strong>the</strong> practical<br />
purposes <strong>of</strong> identification, but this character can be useful when comparing collections micro-<br />
scopically.<br />
A similarly difficult character is <strong>the</strong> relative abundance <strong>of</strong> <strong>the</strong> paraphyses. The two extremes<br />
can be referred to as: (a) 'numerous' - large in number and immediately discernible when<br />
observing a mount in 10% KOH at x400; (b) 'scanty' - few in number and not immediately<br />
obvious when observed in <strong>the</strong> same way. In hymenia with scanty paraphyses <strong>the</strong> 'extra space' is<br />
taken up by hymenial gel or a higher proportion <strong>of</strong> asci, or a combination <strong>of</strong> both. The situation<br />
in most species lies somewhere between <strong>the</strong> two extremes. Fur<strong>the</strong>rmore, <strong>the</strong> relative propor-<br />
tions <strong>of</strong> hymenial gel and asci to paraphyses sometimes increases as <strong>the</strong> apo<strong>the</strong>cium expands (cf<br />
<strong>the</strong> above example <strong>of</strong> M. bauschiana, M. sylvicola, and M. lignaria). An accurate assessment <strong>of</strong><br />
<strong>the</strong> abundance <strong>of</strong> paraphyses is not usually essential for <strong>the</strong> routine identification <strong>of</strong> Micarea<br />
species, although it can be helpful when comparing difficult, convergent forms <strong>of</strong> M. denigrata<br />
and M. misella (see couplet 11 <strong>of</strong> <strong>the</strong> main key).<br />
In addition to <strong>the</strong> 'normal' paraphyses described above, <strong>the</strong> hymenium <strong>of</strong> several species (e.g.<br />
M. bauschiana, M. botryoides, M. eximia, M. nigella, M. sylvicola, andM. tuberculata) contain<br />
scattered individuals, or small fascicles, <strong>of</strong> ra<strong>the</strong>r stout paraphyses. These 'paraphyses' are<br />
about 2-3 /xm wide and usually more distinctly septate than normal paraphyses. Fur<strong>the</strong>rmore<br />
(especially in species with a dark hypo<strong>the</strong>cium), <strong>the</strong>y are <strong>of</strong>ten coated in pigment throughout<br />
<strong>the</strong>ir length (Fig. 34), with <strong>the</strong> result that <strong>the</strong> hymenium is seen to be intersected by dark vertical<br />
streaks. They are mainly found in species with 'scanty' paraphyses, and appear to extend deep<br />
into <strong>the</strong> hypo<strong>the</strong>cium. The elucidation <strong>of</strong> <strong>the</strong>ir true status and function awaits detailed<br />
ontogenetic studies, but it is possible that <strong>the</strong>y have a streng<strong>the</strong>ning, spacing or protective<br />
function during <strong>the</strong> maturation <strong>of</strong> <strong>the</strong> hymenium from <strong>the</strong> primary corpus.<br />
Hypo<strong>the</strong>cium<br />
The area <strong>of</strong> tissue lying below <strong>the</strong> hymenium, or between <strong>the</strong> hymenium and <strong>the</strong> excipulum (if<br />
present), is generally referred to by lichenologists as <strong>the</strong> 'hypo<strong>the</strong>cium'. In many groups <strong>of</strong><br />
lichenized discomycetes (including <strong>the</strong> Lecideaceae) this area can be divided into an upper,<br />
usually narrow, layer containing mainly ascogenous hyphae, and a lower, <strong>of</strong>ten much deeper<br />
layer <strong>of</strong> structural tissue (hyphae gelatinised to various extents, according to genus or species).<br />
These two layers are <strong>of</strong>ten <strong>of</strong> different colour or colour intensity. Where <strong>the</strong> two layers are<br />
.
62 BRIAN JOHN COPPINS<br />
Fig. 34 Two types <strong>of</strong> paraphyses in Micarea tuberculata. A, normal, non-pigmented paraphyses. B, stout,<br />
pigmented paraphyses. Scale = 10 /^m.<br />
distinct <strong>the</strong>y have been referred to as <strong>the</strong> 'subhymenium' and 'hypo<strong>the</strong>cium' respectively (e.g.<br />
Hertel, 19776). The term 'hypo<strong>the</strong>cium' in this context has been substituted by many students <strong>of</strong><br />
non-lichenised discomycetes by <strong>the</strong> term 'medullary excipulum' (Korf, 1973), and was called <strong>the</strong><br />
'ental excipulum' by Eckblad (1968). In Micarea <strong>the</strong> two layers are well-differentiated in only a<br />
few species (e.g. M. crassipes); and in <strong>the</strong> present work 'hypo<strong>the</strong>cium' refers to all ascocarp<br />
tissue lying below <strong>the</strong> hymenium, apart from <strong>the</strong> excipulum (where present). This same<br />
terminology was adopted in <strong>the</strong> recent study <strong>of</strong> Catillaria by Kilias (1981).<br />
The hypo<strong>the</strong>cium in Micarea is composed <strong>of</strong> deeply staining (e.g. in LCB and ammoniacal<br />
erythrosin), swollen-celled ascogenous hyphae (c. 2-5 /am wide) and moderately staining,<br />
slender 'structural' hyphae (c. 0-7-2 fxm wide; slightly varying in width according to species),<br />
embedded in a gelatinous matrix. In species with a colourless or pale hypo<strong>the</strong>cium <strong>the</strong> matrix is<br />
± dispersed in K and <strong>the</strong> hyphae are <strong>the</strong>n clearly visible ( x 400) . In some species with a coloured<br />
hypo<strong>the</strong>cium <strong>the</strong> pigment may be ± evenly distributed through <strong>the</strong> matrix and <strong>the</strong> hypthae are<br />
similarly distinct in K (e.g. M. eximia, M. olivacea, and M. turfosa). However, in <strong>the</strong> majority <strong>of</strong><br />
species with a darkly coloured hypo<strong>the</strong>cium (e.g. M. assimilata, M. botryoides, M. melaena, M.<br />
myriocarpa, M. nigella, M. sylvicola, and M. tuberculata) <strong>the</strong> pigment, although present in <strong>the</strong><br />
matrix, exists as a strongly adhering coat around <strong>the</strong> hyphae, such that <strong>the</strong> hyphae appear<br />
thick-walled and broad (c. 1-5^ /xm). The pigment coatings tend to bind <strong>the</strong> hyphae toge<strong>the</strong>r so<br />
that <strong>the</strong>y are <strong>of</strong>ten indistinct in K. For <strong>the</strong> routine identification <strong>of</strong> Micarea species it is not<br />
essential to know <strong>the</strong> location <strong>of</strong> pigment (i . e . ±<br />
B<br />
H<br />
evenly distributed through <strong>the</strong> gel-matrix versus
LICHEN GENUS MICAREA IN EUROPE 63<br />
tightly bound to hyphae) but such knowledge is sometimes <strong>of</strong> supplementary value, e.g. when<br />
comparing M. eximia versus M. nigella and M. olivacea versus M. tuberculata.<br />
A more important diagnostic character in Micarea is <strong>the</strong> colour <strong>of</strong> <strong>the</strong> hypo<strong>the</strong>cium in water<br />
mounts, and <strong>the</strong> corresponding colour changes obtained by <strong>the</strong> addition <strong>of</strong> KOH and HNO3. A<br />
discussion <strong>of</strong> <strong>the</strong> pigments involved is given under 'chemistry'.<br />
The height <strong>of</strong> <strong>the</strong> hypo<strong>the</strong>cium (in vertical section) is largely dependent on <strong>the</strong> overall size and<br />
(especially in species with a poorly developed excipulum) convexity <strong>of</strong> <strong>the</strong> apo<strong>the</strong>cium. The<br />
measurements given in <strong>the</strong> species descriptions relate to normally developed, non-tuberculate<br />
apo<strong>the</strong>cia. This character is <strong>of</strong>ten very variable for a given species and consequently <strong>of</strong> Uttle<br />
diagnostic value when comparing closely similar species. One exception to this is <strong>the</strong> case <strong>of</strong> M.<br />
contexta (20-90 /am) versus M. melaena (80-160 /xm).<br />
In Lugol's iodine <strong>the</strong> hypo<strong>the</strong>cial tissues are non-amyloid, although <strong>the</strong>re is sometimes a faint<br />
bluing in <strong>the</strong> vicinity <strong>of</strong> ascogeneous hyphae (especially in <strong>the</strong> upper part <strong>of</strong> <strong>the</strong> hypo<strong>the</strong>cium).<br />
Excipulum<br />
The size and distinctiveness <strong>of</strong> <strong>the</strong> excipulum ('ectal excipulum') in Micarea varies greatly<br />
according to species and <strong>the</strong> age <strong>of</strong> <strong>the</strong> apo<strong>the</strong>cium. In species such as M. cinerea, M. crassipes<br />
(Fig. 4B), A/, peliocarpa, and M. ternaria <strong>the</strong> excipulum is sufficiently well developed that <strong>the</strong>ir<br />
young apo<strong>the</strong>cia are <strong>of</strong>ten weakly or distinctly (M. crassipes) marginate in outward appearance.<br />
However, even when well developed and initially distinct, <strong>the</strong> excipulum may become reflexed<br />
and ± occluded as <strong>the</strong> apo<strong>the</strong>cium expands and increases in convexity (Fig. 3A-B) or becomes<br />
tuberculate (Fig. 3C). In many species <strong>the</strong> excipulum is always extremely reduced or absent<br />
(Fig.4A).<br />
When present, <strong>the</strong> excipulum is composed <strong>of</strong> outwardly radiating branched and anastomosing<br />
hyphae that ± separate in K. The hyphae closely resemble paraphyses, but are usually more<br />
dense and more richly branched. With markedly convex apo<strong>the</strong>cia it can be difficult to<br />
distinguish between reflexed portions <strong>of</strong> <strong>the</strong> hymenium and what might be an excipulum. In such<br />
cases <strong>the</strong> excipulum (if present) can be identified in good thin sections by <strong>the</strong> absence <strong>of</strong> asci and<br />
a negative (non-amyloid) reaction to Lugol's iodine. The excipulum <strong>of</strong>ten differs in colour or<br />
colour intensity from <strong>the</strong> hymenium, although a similar colour difference may sometimes be<br />
shown by reflexed parts <strong>of</strong> <strong>the</strong> hymenium.<br />
In M. crassipes and rare forms <strong>of</strong> M. lignaria ('f. gomphillaced') <strong>the</strong> excipular and hypo<strong>the</strong>cial<br />
tissues become vertically extended to form a stipe (Figs 3E, 4B).<br />
Anamorphs (conidial states)<br />
With a few noteworthy exceptions, such as Lindsay (1859, 1872) and Gliick (1899), <strong>the</strong> conidial<br />
states (anamorphs) <strong>of</strong> lichenized fungi have received little detailed attention from taxonomists.<br />
Several recent monographic studies have shown that anamorphs can provide useful characters at<br />
various hierarchial levels <strong>of</strong> classification and <strong>the</strong> reader is referred to Vobis (1980) and Vobis &<br />
Hawksworth (1981) for fur<strong>the</strong>r background information on <strong>the</strong> conidial states <strong>of</strong> lichens.<br />
Within <strong>the</strong> genus Micarea <strong>the</strong>re is a diverse array <strong>of</strong> anamorphic forms, possibly unrivalled by<br />
any o<strong>the</strong>r genus <strong>of</strong> lichens, except perhaps for some <strong>of</strong> <strong>the</strong> genera in <strong>the</strong> Asterothyriaceae<br />
(Vezda, 1979) . Information gained from <strong>the</strong> study <strong>of</strong> anamorphs has proved invaluable to me for<br />
<strong>the</strong> delimitation <strong>of</strong> species in Micarea; indeed, several species frequently occur without<br />
apo<strong>the</strong>cia but with numerous pycnidia, such that a detailed knowledge <strong>of</strong> <strong>the</strong> latter is <strong>of</strong>ten<br />
essential for <strong>the</strong>ir identification (see 'key to species without apo<strong>the</strong>cia').<br />
Conidiomata<br />
The conidiomata are usually pycnidial, and are globose, ovoid, doliiform, or ceriberiform in<br />
shape. They may be immersed (or partly so) within <strong>the</strong> thallus or substratum, sessile, or borne<br />
on stalks (pycnidiophores). When stalked, <strong>the</strong> pycnidia are usually ± doliiform and <strong>the</strong><br />
stalk-tissue is comprised <strong>of</strong> loosely interwoven hyphae bound by a gel matrix which is <strong>of</strong>ten<br />
pigmented. In addition, <strong>the</strong> 'stalk-part' <strong>of</strong>ten includes effete pycnidia (Fig. 35B-D). The stalks<br />
are sometimes branched due to <strong>the</strong> simultaneous development <strong>of</strong> two (or more) pycnidia at <strong>the</strong>
64 BRIAN JOHN COPPINS<br />
Table 1 Conidial states (anamorphs) found in European species <strong>of</strong> Micarea.<br />
Conidium - type Stalked<br />
Micarea micro- meso- macro- pycnidia<br />
adnata<br />
alabastrites<br />
anterior<br />
assimilata<br />
bauschiana<br />
botryoides<br />
cinerea<br />
contexta<br />
crassipes<br />
curvata<br />
denigrata<br />
elachista<br />
eximia<br />
globulosella<br />
hedlundii<br />
incrassata<br />
intrusa<br />
leprosula<br />
lignaria<br />
var. endoleuca<br />
lithinella<br />
lutulata<br />
melaena<br />
melaenida<br />
melaeniza + +<br />
melanobola<br />
misella + + +<br />
muhrii<br />
myriocarpa<br />
nigella<br />
nitschkeana<br />
olivacea<br />
osloensis<br />
peliocarpa<br />
prasina<br />
pycnidiophora + +<br />
rhabdogena<br />
stipitata + +<br />
subleprosula<br />
subnigrata<br />
subviolascens<br />
sylvicola<br />
syno<strong>the</strong>oides<br />
ternaria<br />
tuberculata<br />
turfosa +<br />
Total (species) 25 28<br />
apex <strong>of</strong> stalk tissue (Fig. 35F) or at <strong>the</strong> apex <strong>of</strong> an old pycnidium (Fig. 35D) . Stalked pycnidia are<br />
apparently rare in o<strong>the</strong>r genera <strong>of</strong> crustose lichens (or related fungi). However, I have seen<br />
branched pycnidiophores associated with lignicolous Chaeno<strong>the</strong>copsis spp. growing on <strong>the</strong><br />
moribund thaUi and ascocarps <strong>of</strong> Chaeno<strong>the</strong>ca spp. (several collections in E).<br />
9<br />
9
LICHEN GENUS MICAREA IN EUROPE 65<br />
Fig. 35 Micarea botryoides, conidiomata: diagram showing <strong>the</strong> various means <strong>of</strong> 'stalk' formation and<br />
branching (see text for fur<strong>the</strong>r details). Scale = 100 /xm.<br />
The pycnidia <strong>of</strong> species which consistently lack pigment in <strong>the</strong>ir apo<strong>the</strong>cia (e.g. M. adnata, M.<br />
alabastrites, M. pycnidiophora, and M. stipitata) correspondingly lack pigment in <strong>the</strong>ir wall<br />
tissues. In species with coloured apo<strong>the</strong>cia <strong>the</strong> pycnidial walls are usually (at least in part)<br />
coloured also. For a given species <strong>the</strong> pigment involved is usually that found in <strong>the</strong> hymenium (or<br />
epi<strong>the</strong>cium) <strong>of</strong> <strong>the</strong> apo<strong>the</strong>cia, but in a few cases (e.g. M. lutulata) it is <strong>the</strong> pigment found in <strong>the</strong><br />
hypo<strong>the</strong>cium which is involved. The link between apo<strong>the</strong>cial and pycnidial pigmentation helps<br />
distinguish pycnidia belonging to a Micarea from any intermixed pycnidia belonging to a<br />
non-lichenized or parasitic fungus. It fur<strong>the</strong>r helps to distinguish <strong>the</strong> pycnidia belonging to <strong>the</strong><br />
different species <strong>of</strong> Micarea in mixed populations; for example, <strong>the</strong> stalked pycnidia <strong>of</strong> M.<br />
misella (walls oUvaceous, K+ violet) are <strong>of</strong>ten found intermixed with those <strong>of</strong> o<strong>the</strong>r species such<br />
as M. anterior (walls reddish brown, K— ) and M. melaeniza (walls dark olivaceous, K-).<br />
When a pycnidium is deeply immersed in <strong>the</strong> thallus it is <strong>of</strong>ten only <strong>the</strong> uppermost part around<br />
<strong>the</strong> ostiole which is coloured, and likewise, if <strong>the</strong> pycnidium is semi-immersed it is <strong>of</strong>ten only <strong>the</strong><br />
upper, exposed part which is coloured. With <strong>the</strong> exception <strong>of</strong> <strong>the</strong> species that are characteristically<br />
devoid <strong>of</strong> pigment and some forms <strong>of</strong> o<strong>the</strong>r species in deep shade, <strong>the</strong> walls <strong>of</strong> sessile or<br />
stalked pycnidia are coloured more or less throughout.<br />
As mentioned above, <strong>the</strong> conidiomata <strong>of</strong> Micarea species are pycnidial, but <strong>the</strong>re is one<br />
exception to this. M. adnata has two anamorphic states: one in which <strong>the</strong> conidia (mesoconidia)<br />
are produced internally in immersed pycnidia; and ano<strong>the</strong>r where <strong>the</strong> conidia (macroconidia)<br />
are borne externally on cushion-like sporodochia which resemble small apo<strong>the</strong>cia. This latter<br />
state could be considered to be a hyphomycetous anamorph but comparative ontogenetic<br />
studies are required to investigate <strong>the</strong> <strong>the</strong>oretical possibility that <strong>the</strong>se sporodochia have<br />
evolved from pycnidial conidiomata by an exertion <strong>of</strong> <strong>the</strong> 'hymenium', and a thickening <strong>of</strong> <strong>the</strong><br />
subtending wall-tissue to form a supporting cushion (Fig. 36). Unfortunately this hypo<strong>the</strong>sis is<br />
not supported by intermediate forms in o<strong>the</strong>r species, although <strong>the</strong> pycnidia (with filiform or<br />
curved-hamate macroconidia) <strong>of</strong>, for example, M. cinerea and M. peliocarpa <strong>of</strong>ten have widely
66 BRIAN JOHN COPPINS<br />
Fig. 36 Suggested pathway for evolution <strong>of</strong> a sporodochium (D, as in M. adnata) from a pycnidium (A).<br />
The step A^B can be seen in <strong>the</strong> macroconidial conidiomata <strong>of</strong> e.g. M. cinerea and M. peliocarpa, and<br />
<strong>the</strong> mesoconidial conidiomata <strong>of</strong> e.g. M. denigrata and M. sylvicola. No structure equivalent to 'C is<br />
known from Micarea.<br />
gaping ostioles which eventually expose <strong>the</strong> 'hymenium' . The<br />
<strong>of</strong>ten dubious distinctions<br />
between pycnidia, acervuli, and o<strong>the</strong>r types <strong>of</strong> conidiomata are discussed by Nag Raj (1981).<br />
Although <strong>the</strong> sporodochial anamorph <strong>of</strong> M. adnata has unusually large, oblong-elHpsoid conidia<br />
and <strong>the</strong> longest conidiogenous cells known in <strong>the</strong> genus, <strong>the</strong>re seems to be no fundamental<br />
difference in <strong>the</strong>se features from those in o<strong>the</strong>r Micarea anamorphs, and <strong>the</strong> mode <strong>of</strong><br />
conidiogenesis is apparently <strong>the</strong> same in all cases (see below).<br />
Conidia<br />
Vobis & Hawksworth (1981) estimate that perhaps as many as 8000 species <strong>of</strong> Uchenized fungi<br />
have conidial anamorphs, and that in most cases a given species has only one conidium type.<br />
However, it is becoming increasingly evident that a large number <strong>of</strong> lichens (especially crustose<br />
lichens) have two conidium types, although many <strong>of</strong> <strong>the</strong> examples have not yet been notified in<br />
<strong>the</strong> literature. Some examples from my own studies include Anisomeridium biforme, A.<br />
juistense, Catillaria globulosa, Lecania cyrtellina, Lecanora quercicola, and Opegrapha niveoatra.<br />
Hedlund (1895) found M. denigrata and M. prasina to each have two conidium types and<br />
during <strong>the</strong> present study 16 <strong>of</strong> <strong>the</strong> 45 European species <strong>of</strong> Micarea have been found to have two<br />
conidium types. Even more surprising has been <strong>the</strong> discovery that three species {M. denigrata,<br />
M. nitschkeana, and M. lignaria) each have three conidium types (Figs 42, 45), and to my<br />
knowledge <strong>the</strong>se are <strong>the</strong> first reported instances <strong>of</strong> fungi (whe<strong>the</strong>r lichenized or not) with more<br />
than two pycnidial (coelomycetous) anamorphs. There are, however, a few species <strong>of</strong> sooty
LICHEN GENUS MICAREA IN EUROPE 67<br />
mould (e.g. in <strong>the</strong> Metacapnodiaceae) which have three hyphomycetous anamorphs (Hughes<br />
1972, 1976).<br />
The main distinguishing features <strong>of</strong> <strong>the</strong> three conidium types found in Micarea are as follows<br />
(see Figs 37-52 for example): Microconidia, narrowly cylindrical or narrowly fusiform, aseptate,<br />
eguttulate and not constricted in <strong>the</strong> middle, mostly in <strong>the</strong> range 3-8x0-5-1 /Am; produced in<br />
small immersed or ± sessile pycnidia, mostly c. 20-60 /xm diam. Mesoconidia, variable in shape,<br />
e.g. cylindrical, oblong, ovoid-oblong or obovoid-oblong, aseptate, <strong>of</strong>ten biguttulate and (or)<br />
slightly constricted in <strong>the</strong> middle; produced in small to large, immersed, sessile or stalked<br />
pycnidia. All conidia found in stalked pycnidia are included here. Macroconidia, mostly fihform<br />
or curved and <strong>of</strong>ten septate, or heUcoid and septate (M. subnigrata (p. 183)); produced in<br />
medium-sized to large (<strong>of</strong>ten up to 200 jxm diam or more) pycnidia. Also included here are <strong>the</strong><br />
large, aseptate, eguttulate, oblong-ellipsoid conidia produced by <strong>the</strong> sporodochia <strong>of</strong> M. adnata.<br />
In all cases no single pycnidium has been found to contain more than one conidium type. A<br />
few specimens <strong>of</strong> M. denigrata and M. nitschkeana have been found with all three anamorphs on<br />
<strong>the</strong> same thallus; but such occurrences are rare and it is more usual to find just one or two <strong>of</strong><br />
<strong>the</strong>m. A few species (e.g. M. assimilata, M. crassipes, and M. incrassata appear to have only one<br />
anamorphic state, with conidia somewhat intermediate between microconidia and mesoconidia<br />
as defined above; thus <strong>the</strong> assignment <strong>of</strong> <strong>the</strong>ir conidia to one or o<strong>the</strong>r <strong>of</strong> <strong>the</strong>se types (Table 1)<br />
must be considered tentative. The distinction between <strong>the</strong> three conidium types (especially<br />
micro- and mesoconidia) is most obvious when two or three <strong>of</strong> <strong>the</strong>m occur on <strong>the</strong> same thallus.<br />
The use <strong>of</strong> <strong>the</strong> terms 'micro-', 'meso-' and 'macroconidia' are here apphed solely to <strong>the</strong> conidium<br />
types found in Micarea - <strong>the</strong>y may not necessarily be analogous to <strong>the</strong>ir use in o<strong>the</strong>r lichenized or<br />
non-lichenized fungi. Table 1 indicates <strong>the</strong> conidium type(s) known for each <strong>of</strong> <strong>the</strong> European<br />
species <strong>of</strong> Micarea, and from this a numerical summary <strong>of</strong> <strong>the</strong> various known combinations is as<br />
follows:<br />
Combination <strong>of</strong> conidium types Number <strong>of</strong> species<br />
(anamorphs) (holomorphs)<br />
Micro- -I- meso- -I- macro- 3<br />
Micro- + meso- 10<br />
Micro- -I- macro- 5<br />
Meso- -I- macro- 1<br />
Micro- only 6<br />
Meso- only<br />
Macro- only<br />
14<br />
Anamorph(s) unknown 6<br />
The role <strong>of</strong> each <strong>of</strong> <strong>the</strong> conidium types is as yet unknown. Vobis (1977) obtained successful<br />
germination and subsequent growth <strong>of</strong> mycelium from <strong>the</strong> macroconidia <strong>of</strong> Lecanactis abietina,<br />
but was unable to germinate <strong>the</strong> microconidia <strong>of</strong> <strong>the</strong> same species. His results suggest that <strong>the</strong><br />
former may well act as asexual propagules, and <strong>the</strong> latter as spermatia in sexual reproduction. It<br />
seems most likely to me that <strong>the</strong> microconidia oi Micarea species are spermatia. As to whe<strong>the</strong>r or<br />
not <strong>the</strong>y are essential components <strong>of</strong> <strong>the</strong> sexual reproductive process is much less certain; 21 <strong>of</strong><br />
<strong>the</strong> treated species are not known to produce microconidia.<br />
The role <strong>of</strong> conidia as asexual propagules is still a matter <strong>of</strong> much debate and speculation<br />
among lichenologists, but such a role (at least in certain cases) can be inferred from <strong>the</strong> fact that a<br />
few crustose species (e.g. Lecanactis subabietina and an undescribed Bacidia) are not known to<br />
have apo<strong>the</strong>cia or vegetative diaspores (i.e. soredia, isidia, etc.) but always have numerous<br />
pycnidia. To such examples can be added <strong>the</strong> numerous crustose lichens whose apo<strong>the</strong>cia are<br />
known, but which commonly occur as sterile populations with numerous pycnidia, e.g.<br />
Anisomeridium juistense (macroconidial state), Arthonia phaeobaea, A. spadicea, Bacidia<br />
arnoldiana, B. carneoglauca, B. trachona, Cliostomum graniforme, C. griffithii, Lecanactis<br />
abietina, Lecidea erratica, Opegrapha niveoatra, O. vermicellifera, and O. vulgata. This last<br />
group also includes Micarea botryaides, M. denigrata, M. misella, M. pycnidiophora, and M.<br />
stipitata, all <strong>of</strong> which commonly occur with abundant mesoconidia-containing pycnidia and few<br />
(if any) mature apo<strong>the</strong>cia.
68 BRIAN JOHN COPPINS<br />
Fig. 37 M. adnata (E - holotype), macroconidia and conidiogenous cells. Scale = 10 />tm.
LICHEN GENUS MICAREA IN EUROPE 69<br />
00 B<br />
Fig. 38 A, M. botryaides (Coppins 8429, E), mesoconidia and conidiogenous cells. B, M. melaeniza (S -<br />
holotype), mesoconidia. C-D, M. anterior Sweden, Angermanland, Langsele, 1892, Hedlund, S); C,<br />
mesoconidia; D, microconidia. Scale = 10 yam.
70 BRIAN JOHN COPPINS<br />
s<br />
o<br />
o<br />
c<br />
o<br />
c<br />
o<br />
-o cCO<br />
u<br />
6<br />
I<br />
ei)
\r\ r\<br />
[J^\J<br />
LICHEN GENUS MICAREA IN EUROPE 71<br />
r\<br />
\J<br />
Fig. 40 A-B, M. contexta (S - lectotype); A, mesoconidia; B, microconidia. C, M. eximia (Malme Lich.<br />
suec. exs. 26, C), mesoconidia. D, M. bauschiana {Coppins 4111, E), microconidia and conidiogenous<br />
cells. Scale = lOjum.
72 BRIAN JOHN COPPINS<br />
n<br />
1/<br />
[\ f] n<br />
Fig. 41 A-B, M. incrassata, microconidia; A, (S - holotype); B, (Kerguelen, 1875, Eaton, E). C, M.<br />
crassipes (Vezda Lich. Sel. 11, BM), (?)mesoconidia and conidiogenous cells. Scale = 10 /tm.<br />
B
LICHEN GENUS MICAREA IN EUROPE 73<br />
Fig. 42 M. denigrata (Coppins 1888, E), conidia with conidiogenous cells; A, macroconidia; B, mesoconi-<br />
dia; C, microconidia. Scale = 10 /Am.
74 BRIAN JOHN COPPINS<br />
Fig. 43 A-B, M. globulosella (Czechoslovakia, Slovakia, Vysoke Tatry, 1879, Lojka, BM); A, mesoconidia;<br />
B, microconidia. C, M. globulosella (S - lectotype), mesoconidia. D-E, M. syno<strong>the</strong>oides (Coppins<br />
2942, E); D, mesoconidia; E, microconidia. Scale = 10 /jun.
LICHEN GENUS MICAREA IN EUROPE 75<br />
Fig. 44 A, M. hedlundii (UPS - holotype), mesoconidia and conidiogenous cells. B, M. lutulata (Wales,<br />
Pembroke, Tycanol, 1980, James, BM), mesoconidia and conidiogenous cells. Scale = 10 )Ltm.
76 BRIAN JOHN COPPINS<br />
Fig. 45 M. lignaria. A, macroconidia (Coppins 4658, E).<br />
microconidia (Coppins 8952, E). Scale = 10 ixm.<br />
B<br />
B, mesoconidia (Coppins 8436, E). C,
LICHEN GENUS MICAREA IN EUROPE 77<br />
\j \l<br />
Fig. 46 A, M. melaena {Coppins 6041, E), macroconidia. B-C, M. melanobola (H-NYL 2164<br />
lectotype); B, mesoconidia; C, microconidia. Scale = 10/i,m.<br />
B
78 BRIAN JOHN COPPINS<br />
B<br />
OOQog<br />
Fig. 47 A, M. olivacea (BM - holotype), mesoconidia and conidiogenous cells. B, M. nigelta (E -<br />
holotype), mesoconidia and conidiogenous cells. C, M. myriocarpa (Coppins 8939, E), mesoconidia and<br />
conidiogenous cells. Scale = 10 /xm.
LICHEN GENUS MICAREA IN EUROPE 79<br />
Fig. 48 M. peliocarpa (England, New Forest, Great Wood, Bramble Hill Walk. 1970, Coppins et al., E).<br />
A, macroconidia and conidiogenous cells. B, microconidia and conidiogenous cells. Scale = 10 jxm.
80 BRIAN JOHN COPPINS<br />
Fig. 49 A-B, M. prasina (GZU - holotype <strong>of</strong> M. polytrichi); A, mesoconidia; B, microconidia. C, M.<br />
prasina (Coppins 8009, E), mesoconidia. D, M. prasina {Coppins 2835, E), microconidia. Scale — 10<br />
fim.<br />
{\<br />
^<br />
B
LICHEN GENUS MICAREA IN EUROPE 81<br />
Fig. 50 M. subnigrata (Coppins 8417, E). A, macroconidia and conidiogenous cells. B, microconidia and<br />
conidiogenous cells. Scale = 10 /u,m.<br />
B
82 BRIAN JOHN COPPINS<br />
Fig. 51 A, M. sylvicola {Muhr 2579, E), mesoconidia and conidiogenous cells. B, M. tuberculata (O -<br />
lectotype), mesoconidia and conidiogenous cells. Scale = 10 /x,m.<br />
B
LICHEN GENUS MICAREA IN EUROPE 83<br />
Fig. 52 A-B, M. ternaria, mesoconidia; A, (H-NYL 18682 - holotype); B, {Thomson 9188, DUKE). C,<br />
M. aff. ternaria (Fair Isle, 1976, Duncan, BM), mesoconidia. Scale = 10 fjun.<br />
B
84<br />
BRIAN JOHN COPPINS<br />
In <strong>the</strong> case <strong>of</strong> M. denigrata it is a general rule (but with exceptions) that when on worked wood<br />
(fence-posts, garden furniture, window frames, etc.) its thallus has numerous pycnidia (with<br />
mesoconidia) and <strong>of</strong>ten few (if any) mature apo<strong>the</strong>cia. On natural substrata (e.g. fallen tree<br />
trunks in old woodlands) pycnidia with microconidia or macroconidia are more prevalent,<br />
although <strong>the</strong>y are usually relatively fewer in number and associated with numerous apo<strong>the</strong>cia. It<br />
seems highly likely that <strong>the</strong> success <strong>of</strong> M. denigrata as a primary coloniser <strong>of</strong> newly available<br />
substrata is largely due to <strong>the</strong> successful role <strong>of</strong> its mesoconidia as asexual propagules. Its<br />
microconidia probably function as spermatia, but <strong>the</strong> function <strong>of</strong> <strong>the</strong> macroconidia is less<br />
obvious, although I suspect that <strong>the</strong>y, like <strong>the</strong> mesoconidia, act as asexual diaspores (perhaps in<br />
a different way).<br />
Ano<strong>the</strong>r observation pertinent to this discussion is that I have frequently observed <strong>the</strong> mesoand<br />
macroconidia <strong>of</strong> Micarea species being extruded through <strong>the</strong> ostioles <strong>of</strong> <strong>the</strong> pycnidia as<br />
white mucilaginous blobs; such phenomena are usually seen after periods <strong>of</strong> wet wea<strong>the</strong>r. It is<br />
tempting to suggest that <strong>the</strong>se extrusions facilitate dispersal beyond <strong>the</strong> limits <strong>of</strong> <strong>the</strong> parent<br />
thallus by <strong>the</strong> action <strong>of</strong> rain or passing arthropods and mollusca. Dispersal by invertebrates may<br />
be <strong>of</strong> especial importance for species such as M. botryoides, which occur in sheltered situations<br />
rarely subjected to rain-wash. I have only rarely observed microconidia being extruded as white<br />
blobs, thus suggesting that <strong>the</strong>re is no need for <strong>the</strong>m to be dispersed beyond <strong>the</strong> limits <strong>of</strong> <strong>the</strong><br />
parent thallus. Indeed, if microconidia do function as spermatia, and if <strong>the</strong> mycobiont <strong>of</strong> <strong>the</strong><br />
Micarea is homothallic, <strong>the</strong>re would be no requirement for <strong>the</strong>m to be dispersed more than a few<br />
millimetres.<br />
Conidiogenous cells<br />
The conidiogenous cells <strong>of</strong> Micarea species are always phialidic, although in many cases <strong>the</strong><br />
phialides are seen to undergo 'percurrent proliferation' (Figs 42B, 44B). The conidiogenous<br />
cells belong to Types I or II <strong>of</strong> Vobis & Hawksworth (1981) and arise from <strong>the</strong> inner wall <strong>of</strong> <strong>the</strong><br />
pycnidium (or on <strong>the</strong> outer surface <strong>of</strong> <strong>the</strong> sporodochium in M. adnata). Their subtending cells<br />
are never sufficiently regular in shape for <strong>the</strong>m to be termed 'conidiophores'. In several species<br />
whose pycnidial walls are intensely pigmented <strong>the</strong> bases <strong>of</strong> <strong>the</strong> conidiogenous cells are <strong>of</strong>ten<br />
similarly pigmented (Figs 47B, 51B). The nearest approach to <strong>the</strong> differentiation <strong>of</strong> wall-tissue<br />
from a conidiogenous layer is found in <strong>the</strong> thick-walled pycnidia <strong>of</strong> M. elachista.<br />
There is much variety in <strong>the</strong> shape <strong>of</strong> conidigenous cells (e.g. ampuUiform, doliiform,<br />
lageniform, cyUndrical) and <strong>the</strong>re is sometimes much variation within a single pycnidium<br />
(Figs 42B, 44A). Conidiogenous cells with long cyhndrical necks <strong>of</strong>ten have swollen bases (Figs<br />
38A, 47B, 5 IB). Although critical observations and measurements have not been made for all<br />
species, <strong>the</strong> size and shape <strong>of</strong> conidiogenous cells have rarely been found to be useful characters<br />
for <strong>the</strong> separation <strong>of</strong> closely similar species. However, one exceptional case is that <strong>of</strong> saxicolous<br />
forms <strong>of</strong> M. olivacea (Fig. 47A) versus M. tuberculata (Fig. 5 IB), in which <strong>the</strong> conidiogenous<br />
cells <strong>of</strong> <strong>the</strong> latter are much longer.<br />
Chemistry<br />
The discussions below are presented under two sub-headings. The first deals with 'lichen<br />
substances', which are readily extractable in acetone and identifiable by thin-layer chromatography<br />
(t.l.c; see 'Methods'). The second part deals with pigments that cannot be analysed in this<br />
way; <strong>the</strong>ir chemical nature is at present unknown and <strong>the</strong>y can only be characterised by <strong>the</strong>ir<br />
colour in water and subsequent reactions with reagents such as K and HNO3.<br />
Lichen substances<br />
Prior to <strong>the</strong> present studies <strong>the</strong>re is little evidence in <strong>the</strong> literature to suggest that species <strong>of</strong><br />
Micarea contain any lichen substances. However, <strong>the</strong>re are a few hints given in some early<br />
descriptions: for example, Leighton (1879: 362) gives 'K+ yellow, C+ orange-red' reactions for<br />
Lecidea milliaria [Micarea lignaria], which probably relate to his specimens <strong>of</strong> <strong>the</strong> var.<br />
endoleuca. The only reported t.l.c. analysis <strong>of</strong> a Micarea appears to be that <strong>of</strong> Huneck &
LICHEN GENUS MICAREA IN EUROPE 85<br />
Table 2 Chemical content and corresponding spot test reactions <strong>of</strong> European species <strong>of</strong> Micarea known to<br />
contain lichen substances. +, present; ±, presence variable and sometimes absent; ?, present occa-<br />
sionally as trace amounts or contaminant; R, red; Y, yellow; fY, faint yellow; O, persistent orange; ( )<br />
reactions not obtainable in many collections; *, including 3 or 4 accessory substances.
86 BRIAN JOHN COPPINS<br />
6 -<br />
5h<br />
4<br />
3<br />
2<br />
1<br />
r m^<br />
B<br />
C<br />
F F<br />
D D<br />
Fig. 53 Diagram <strong>of</strong> chromatogram in solvent system TDA.<br />
^<br />
K<br />
J<br />
(22) (^<br />
8<br />
TDA<br />
1, Control (Parmelia acetabulum plus Cladonia subcervicornis). 2, M. peliocarpa. 3, M. subleprosula. 4,<br />
M. lignaria. 5, M. leprosula. 6, M. lignaria var. endoleuca. 7, M. prasinas. lat. 8, M. prasinas. str. 9, M.<br />
prasina s. lat. A, atranorin, B, norstictic acid. C, fumarprotocetraric acid. D, gyrophoric acid. E,<br />
alectorialic acid, and accessory substances (E\ E^). F, argopsin. G and H, xanthones. J, prasina<br />
unknown A. K, prasina unknown B. L, prasina unknown C.<br />
7 r<br />
6<br />
5<br />
4<br />
3<br />
2<br />
1<br />
B<br />
(Si<br />
D
LICHEN GENUS MICAREA IN EUROPE 87<br />
most widely occurring compounds among lichen genera. It has been found in 11 Micarea species,<br />
although its presence in M. curvata has only been inferred from spot tests, <strong>the</strong> solitary specimen<br />
(holotype) being too small for t.l.c. analysis. In addition to <strong>the</strong>se occurrences, gyrophoric acid is<br />
frequently detected in trace amounts in chromatograms <strong>of</strong> o<strong>the</strong>r taxa, especially M. prasina s.<br />
ampl. The fact that Micarea species <strong>of</strong>ten grow intermixed with o<strong>the</strong>r lichens, including several<br />
that contain gyrophoric acid (e.g. Lecidea icmalea and L. granulosa agg.) raises <strong>the</strong> question as<br />
to whe<strong>the</strong>r <strong>the</strong>se trace amounts represent its presence as an accessory substance or as a<br />
contaminant. This is difficult to answer. Gyrophoric acid is certainly found in <strong>the</strong> M. prasina<br />
complex, as it has been detected in large amounts in <strong>the</strong> type material <strong>of</strong> Lecidea levicula Nyl.<br />
from Cuba. (Fur<strong>the</strong>r studies are required to establish <strong>the</strong> taxonomic status <strong>of</strong> L. levicula.)<br />
Well-developed specimens <strong>of</strong> M. denigrata, M. nitschkeana, and M. melaena have an areolate<br />
thallus containing readily detectable quantities <strong>of</strong> gyrophoric acid. However, <strong>the</strong> thallus <strong>of</strong> <strong>the</strong>se<br />
species is sometimes scurfy-granular and blackish due to disruption by invading dematiaceous<br />
fungi and non-lichenized algae, and gyrophoric acid is produced in very low amounts or is<br />
apparently absent altoge<strong>the</strong>r (not detectable by t.l.c).<br />
The most surprising result <strong>of</strong> <strong>the</strong> chemical studies in Micarea was <strong>the</strong> discovery <strong>of</strong> three<br />
unknown, but very distinctive, compounds in M. prasina. The three compounds can be<br />
characterised thus (UV at 254 m/x):<br />
prasina unknown A. TDA 5: HEF 6, UV-h blue-white. After H2SO4 and charring: UV+ dull<br />
orange-red, dull orange in daylight.<br />
prasina unknown B. TDA 5: HEF 6, UV+ blue (less bright than 'unknown A'). After H2SO4<br />
and charring: UV-f- vivid citrine-yellow, yellow (without orange tinge) in daylight.<br />
prasina unknown C. TDA 4-5: HEF 6-7 (slightly higher than 'unknowns A and B'), UV-I-<br />
grey or pale mauve (TDA) or ± colourless (HEF). After H2SO4 and charring: UV-I- violet-blue,<br />
± colourless in daylight but turning a pale pinkish-lilac after several weeks.<br />
In all <strong>the</strong> numerous European and North American specimens <strong>of</strong> M. prasina s. ampl.<br />
examined so far, <strong>the</strong>se compounds have never been found in combination, and M. prasina exists<br />
as three distinctive chemical races (see taxonomic account <strong>of</strong> M. prasina for fur<strong>the</strong>r discussion).<br />
None <strong>of</strong> <strong>the</strong> three compounds can be identified in <strong>the</strong> tabulations <strong>of</strong> Culberson (1972), and <strong>the</strong>y<br />
have never been encountered in o<strong>the</strong>r genera during <strong>the</strong> numerous and diverse investigations by<br />
Mr P. W. James (pers. comm.). Samples <strong>of</strong> specimens containing 'unknowns A and B' are<br />
currently being studied by Dr J. A. Elix.<br />
The identification <strong>of</strong> hchen substances (even if only by spot tests) is essential for distinguishing<br />
sterile specimens <strong>of</strong> M. leprosula from M. subleprosula, and for separating <strong>the</strong> two varieties <strong>of</strong><br />
M. lignaria; it is also <strong>of</strong> great value in <strong>the</strong> routine identification <strong>of</strong> many o<strong>the</strong>r species (see 'Keys<br />
to species'). The three European races <strong>of</strong> M. prasina are not distinguishable by spot tests, but are<br />
readily identified by t.l.c.<br />
Pigments<br />
The nature <strong>of</strong> pigments and <strong>the</strong>ir location (especially within apo<strong>the</strong>cia) are <strong>of</strong> prime importance<br />
in <strong>the</strong> delimitation <strong>of</strong> species within many lichen genera, particularly those included in <strong>the</strong><br />
Lecideaceae, and Micarea is no exception. Little is known <strong>of</strong> <strong>the</strong>se acetone insoluble pigments<br />
and a detailed knowledge <strong>of</strong> <strong>the</strong>ir structure and biogenesis would be invaluable for <strong>the</strong><br />
evaluation <strong>of</strong> <strong>the</strong>ir taxonomic significance.<br />
On <strong>the</strong> basis <strong>of</strong> <strong>the</strong> colours seen in water mounts and <strong>the</strong> colour changes brought about by <strong>the</strong><br />
application <strong>of</strong> a strong alkali (KOH) and a strong acid (HNO3) at least eight pigments can be<br />
recognized in Micarea. A very provisional summary <strong>of</strong> <strong>the</strong>se pigments is given below:<br />
Pigment A. Green or aeruginose, K- or -I- green intensifying, HNO3+ red; in various tissues (according<br />
to species) <strong>of</strong> M. assimilata, M. bauschiana, M. cinerea, M. intrusa, M. lignaria, M. melaena, M.<br />
peliocarpa, M. sylvicola, and M. tuberculata.<br />
Pigment B. Purple, K+ green, HNO3+ purple-red; mostly in <strong>the</strong> hypo<strong>the</strong>cium <strong>of</strong> M. assimilata, M.<br />
contexta, M. crassipes, M. eximia, M. melaena, M. nigella, and M. sylvicola (rare forms).<br />
Pigment C. Purple, K+ purple intensifying (sometimes partly dissolving into solution); occasionally in
.<br />
88 BRIAN JOHN COPPINS<br />
<strong>the</strong> epi<strong>the</strong>cium (M. melaenida and forms <strong>of</strong> M. crassipes and M. melaend) but mainly in <strong>the</strong> hypo<strong>the</strong>cium<br />
{M. assimilata, M. crassipes, M. melaena, M. melaenida, and M. subviolascens)<br />
Pigment D. Olivaceous, K+ violet (also C+ violet), HNO3+ red; in various tissues (but rarely in <strong>the</strong><br />
hypo<strong>the</strong>cium, and <strong>the</strong>n in low concentrations) <strong>of</strong> M. denigrata, M. elachista (mostly in pycnidia), M.<br />
globulosella, M. hedlundii, M. melanobola, M. misella, M. nitschkeana, M. prasina, M. subviolascens, and<br />
M. syno<strong>the</strong>oides<br />
Pigment E. Fuscous brown, K+ dissolving into solution, HNO3—; in <strong>the</strong> epi<strong>the</strong>cia <strong>of</strong> M. elachista SiXxdM.<br />
rhabdogena.<br />
Pigment F. Brown (sometimes slightly tinged reddish), K- (not dissolving), HNO3- or + orangebrown;<br />
in various tissues (according to species) <strong>of</strong> A/, botryoides, M. curvata, M. incrassata, M. lutulata, M.<br />
muhrii, M. myriocarpa, M. osloensis, M. subnigrata, andM. turfosa. It is quite possible that more than one<br />
pigment is involved here.<br />
Pigment G. Dilute yellowish, K+ purple (oily droplets), HNO3- ; in goniocysts and sometimes <strong>the</strong> lower<br />
hymenium and hypo<strong>the</strong>cium <strong>of</strong> M. hedlundii.<br />
Pigment H. Pale yellowish orange, K+ purple, HNO3— ; in cytoplasm <strong>of</strong> some ascogenous hyphae, asci,<br />
and spores <strong>of</strong> M. intrusa. This is possibly similar (or identical) to pigment G.<br />
Pigment complexes involving mixtures <strong>of</strong> pigments A, B, and C are <strong>of</strong>ten found (especially in<br />
<strong>the</strong> hypo<strong>the</strong>cium) in M. assimilata, M. contexta, M. crassipes, M. melaena, M. subviolascens,<br />
and M. sylvicola; see <strong>the</strong> individual species accounts for fur<strong>the</strong>r details. The existence <strong>of</strong> such<br />
complexes suggests that pigments A, B, and C are closely related chemically.<br />
Pigment A is probably identical to that found (usually in <strong>the</strong> epi<strong>the</strong>cium) in a large number <strong>of</strong><br />
lichens, especially in Led^efl 5. lat., Catillarias. lat., Bacidias. lat. and Lecanoras. lat. Pigment<br />
C is probably <strong>of</strong> rare occurrence outside Micarea, but I know <strong>of</strong> it in <strong>the</strong> epi<strong>the</strong>cium <strong>of</strong> Bacidia<br />
beckhausii and in several species <strong>of</strong> Pertusaria (e.g. P. hymenea and P. oculata). I have not<br />
encountered pigment E in any o<strong>the</strong>r lichens, but it should be compared with <strong>the</strong> pigment(s)<br />
responsible for <strong>the</strong> ionomidotic reaction found in several genera <strong>of</strong> non-lichenized discomycetes<br />
(Korf , 1973). Pigment F is probably <strong>of</strong> wide occurrence, but <strong>the</strong>re may be several pigments that<br />
impart a brown or 'melanized' appearance in tissues.<br />
The dark violet-blue, K-h aeruginose granules occasionally seen in <strong>the</strong> hymenium <strong>of</strong> M.<br />
contexta and M. lignaria (and also <strong>the</strong> apo<strong>the</strong>cia <strong>of</strong> Lecidea hypnorum and Dactylospora<br />
lobariella), is probably related to (if not <strong>the</strong> same as) pigment B, and it may be <strong>the</strong> same as <strong>the</strong><br />
epi<strong>the</strong>cial pigment found in Bacidia absistens, Mycoblastus fucatus , and Schaereria tenebrosa.<br />
Ecology<br />
No detailed ecological studies have been made in connection with this primarily taxonomic study<br />
<strong>of</strong> Micarea, and much <strong>of</strong> <strong>the</strong> discussion given below is based on floristic notes and casual field<br />
observations. For most amateur and pr<strong>of</strong>essional lichenologists alike, <strong>the</strong> species oi Micarea are<br />
little known, poorly understood, and much overlooked in <strong>the</strong> field. For this reason <strong>the</strong> following<br />
account is more <strong>of</strong> a guide to collectors, ra<strong>the</strong>r than an ecological dissertation. Additional<br />
ecological notes are also included in <strong>the</strong> taxonomic accounts <strong>of</strong> each <strong>of</strong> <strong>the</strong> Micarea species<br />
treated. Syntaxonomic nomenclature follows James ef a/. (1977).<br />
General habitats<br />
Micarea species occur in a wide range <strong>of</strong> habitats but are confined to substrata with a low pH<br />
(below c. pH 5) and generally avoid nutrient enriched situations. Thus, <strong>the</strong>y are absent from<br />
limestone rocks, tops <strong>of</strong> bird-perching stones, seashores, and basic bark. There are a few partial<br />
exceptions, for example M. denigrata can occur on wooden fencing and o<strong>the</strong>r timber in<br />
farmyards, M. prasina can occur on soil and debris among rocks in <strong>the</strong> upper seashore, and M.<br />
lignaria has been found on exposed limestone growing over mats or cushions <strong>of</strong> moribund<br />
bryophytes from which free calcium ions have presumably been leached out. The bark <strong>of</strong> Acer<br />
and Ulmus normally has a pH which is too high for species <strong>of</strong> Micarea (with <strong>the</strong> exception <strong>of</strong> M.<br />
prasina in some situations), but in areas heavily polluted by sulphur dioxide and its derivatives<br />
(e.g. West Yorkshire Conurbation), bark pH can be substantially lowered (Gilbert, 1970), thus<br />
providing a suitable substratum for species such as M. melaena (q.v.) and M. botryoides (q.v.).<br />
.
LICHEN GENUS MICAREA IN EUROPE 89<br />
The principal habitats (I-XI) in which Micarea spp. are found are given below; each is<br />
provided with a list <strong>of</strong> <strong>the</strong> species which can be expected to occur in it (allowing for climatic and<br />
phytogeographical variations). Species given in square brackets [ ] are rarely and unusually<br />
found in <strong>the</strong> given habitat, and those prefixed with a dagger (t) are not known in this habitat<br />
from <strong>the</strong> <strong>British</strong> Isles.<br />
I.<br />
Sheltered, dry underhangs, usually in valley woodland or narrow ravines, including dry undersides<br />
<strong>of</strong> up-ended tree root systems in woodland; growing on rock, loose stones, consolidated soil, and<br />
exposed roots.<br />
M. bauschiana M. myriocarpa<br />
M. botryoides M. sylvicola<br />
M. lignaria M. tuberculata<br />
M. lutulata<br />
II. Exposed parts <strong>of</strong> sheltered rocks, usually in woodland; more frequently wetted than I; growing<br />
directly on rock.<br />
M. botryoides M. sylvicola<br />
tM. curvata [M. cinerea]<br />
M. lignaria s. lat. [M. denigrataj<br />
M. lithinella '\IM. muhrii]<br />
M. melaena [M. nitschkeana]^<br />
M. olivacea [M. prasina]<br />
M. peliocarpa<br />
a, stones amongst Calluna in heathland.<br />
III. Over bryophytes on rocks, boulders, old stumps, or fallen trees in woodland at low altitudes (500 m<br />
in <strong>the</strong> <strong>British</strong> Isles).<br />
M. adnata M. lignaria s. lat.<br />
M. botryoides M. melaena<br />
M. cinerea M. peliocarpa<br />
M. leprosula M. prasina<br />
[M. stipitata]<br />
IV. Exposed, hard siliceous rocks; growing directly on rock.<br />
M. intrusa tM. subviolascens<br />
M. lignaria M. aff . ternaria^<br />
M. subnigrata<br />
b, coastal districts only.<br />
V. Over bryophytes or peaty debris on exposed turf or on, or amongst, rocks and boulders in open<br />
situations in upland, montane, or 'arctic' districts.<br />
M. assimilata M. peliocarpa<br />
tM. crassipes M. subleprosula<br />
M. incrassata tM. ternaria<br />
M. leprosula M. turfosa<br />
M. lignaria s. lat. [M. cinerea]<br />
M. melaena<br />
VI. Over bryophytes or plant debris on <strong>the</strong> ground in old dunes, disused lead and zinc mines, sea-cliffs,<br />
or by woodland tracks.<br />
M. botryoides M. prasina<br />
M. leprosula [M. denigrataj<br />
M. lignaria s. str. flM. misella]<br />
M. peliocarpa<br />
VII. On bare mineral soil at low altitudes.<br />
tM. melaenida [M. lignaria]<br />
tM. osloensis [M. prasina]<br />
[M. leprosula]<br />
VIII. Corticate trunks <strong>of</strong> healthy trees; usually in woodland; sometimes overgrowing bryophytes (*).<br />
*M. alabastrites *M. pycnidiophora<br />
*M. cinerea *M. stipitata
90 BRIAN JOHN COPPINS<br />
fM. elachista *M. syno<strong>the</strong>oides<br />
tM. globulosella [M. botryoides]<br />
M. melaena [M. denigrata]<br />
tM. melanobola [M. leprosula]<br />
*M. peliocarpa [M. lignaria]<br />
*M. prasina [M. nitschkeana]<br />
IX. On attached twigs <strong>of</strong> trees or large shrubs, or thin stems <strong>of</strong> small shrubs; in woodland, heathland, or<br />
scrub.<br />
tM. cinerea M. peliocarpa<br />
M. lignaria s. str. M. prasina<br />
M. nitschkeana [M. denigrata]<br />
X. On timber (i.e. worked wood), e.g. fencing, garden furniture, old window frames, and shingles.<br />
M. cinerea M. nitschkeana<br />
M. denigrata M. peliocarpa<br />
tM. elachista [M. globulosella]<br />
M. lignaria s. str. [M. leprosula]<br />
M. melaena [M. sylvicola]<br />
M. misella<br />
XI. Directly on lignum <strong>of</strong> old stumps and decorticate trunks.<br />
M. adnata M. misella<br />
M. alabastrites tM. muhrii<br />
tM. anterior M. nigella<br />
M. cinerea M. nitschkeana<br />
tM. contexta M. olivacea<br />
M. denigrata M. peliocarpa<br />
tM. elachista M. prasina<br />
tM. eximia tM. rhabdogena<br />
tM. hedlundii /M. botryoides]<br />
M. lignaria s. lat. [M. leprosula]<br />
M. melaena [M. myriocarpa]<br />
tM. melaeniza [M. syno<strong>the</strong>oides]<br />
From <strong>the</strong> above lists it will be seen that several species (e.g. M. lignaria, M. melaena, M.<br />
peliocarpa, and M. prasina) inhabit a very wide range <strong>of</strong> habitats and substrate. At <strong>the</strong> o<strong>the</strong>r<br />
extreme <strong>the</strong>re are many species which are much more restricted e.g. M. tuberculata (I), M.<br />
subnigrata (IV), M. assimilata (V), M. melaenida (VIII), M. pycnidiophora (VIII), and M.<br />
anterior, M. contexta, M. eximia, and M. melaeniza (XI).<br />
In most <strong>of</strong> <strong>the</strong> communities in which <strong>the</strong>y occur Micarea species are usually <strong>of</strong> minor<br />
importance with regard to cover values. However, <strong>the</strong>re are some exceptions such as <strong>the</strong><br />
Micareetum sylvicolae association <strong>of</strong> underhangs and exposed tree root-system (see below), <strong>the</strong><br />
community dominated by M. prasina on trunks in dense conifer plantations, some lignicolous<br />
assemblages on fallen trunks, old stumps, and worked wood. Micarea nitschkeana on Calluna<br />
twigs and litter in some lowland heaths, M. melaena on sandy or peaty soil in some heathlands<br />
and moorlands, and M. lignaria on <strong>the</strong> ground in some old lead mine workings.<br />
Specific habitats<br />
Deciduous (broad-leaved) woodland<br />
In Britain <strong>the</strong> genus JWicarea is best represented in terms <strong>of</strong> number <strong>of</strong> species per site in <strong>the</strong><br />
mature, ± natural woodland (both deciduous and coniferous) on acid soils, in areas with a high<br />
annual rainfall (at least 1000 mm distributed over at least 160 'wet days'; see Coppins, 1976). The<br />
best examples <strong>of</strong> 'Mcarefl-rich' deciduous (broad-leaved) woodlands are found in Wales, <strong>the</strong><br />
English Lake District and <strong>the</strong> west <strong>of</strong> Scotland north <strong>of</strong> <strong>the</strong> Clyde Estuary. The Micarea species<br />
found in <strong>the</strong>m on trees and stumps are included in lists III, VII, and XI above, except that M.<br />
misella, M. nigella, and M. olivacea are primarily species <strong>of</strong> coniferous woodlands.<br />
The occurrence <strong>of</strong> Micarea spp. on bark (or over bryophytes <strong>the</strong>reon) is favoured by leaching,<br />
and is fur<strong>the</strong>r favoured, to <strong>the</strong> detriment <strong>of</strong> more basicolous lichens, by <strong>the</strong> effects <strong>of</strong> 'acid rain'
LICHEN GENUS MICAREA IN EUROPE 91<br />
(Anon, 1980; Fowler et al., 1982). Acid rain effects, resulting from <strong>the</strong> pollution emitted from<br />
<strong>the</strong> industrial areas <strong>of</strong> west-central Scotland (Clydeside and Glasgow conurbation), are probably<br />
<strong>the</strong> explanation for <strong>the</strong> prevalence on <strong>the</strong> trunks <strong>of</strong> mature trees (e.g. Quercus, Alnus,<br />
Betula, and Fraxinus) <strong>of</strong> such species as M. alabastrites, M. cinerea, M. peliocarpa, M. stipitata,<br />
and M. syno<strong>the</strong>oides in <strong>the</strong> deciduous woodlands <strong>of</strong> <strong>the</strong> Cowal Peninsula in sou<strong>the</strong>rn Argyllshire.<br />
The communities in which <strong>the</strong>se species occur are probably referable to <strong>the</strong> Parmelietum<br />
laevigatae, and in areas subjected to 'acid rain' it appears that <strong>the</strong>se communities have to some<br />
extent, and in certain situations (especially on Quercus and Fraxinus), replaced <strong>the</strong> more<br />
basicolous communities <strong>of</strong> <strong>the</strong> Lobarion pulmonariae.<br />
In <strong>the</strong> deciduous woodlands <strong>of</strong> drier parts <strong>of</strong> <strong>the</strong> <strong>British</strong> Isles (annual rainfall <strong>of</strong>
92 BRIAN JOHN COPPINS<br />
Saxicolous habitats<br />
Of <strong>the</strong> 45 European species <strong>of</strong> Micarea, 21 have been found growing directly on rock (lists I,<br />
II, and IV), although <strong>the</strong> genus is poorly represented in <strong>the</strong> main saxicolous lichen alliances <strong>of</strong><br />
exposed siliceous rocks (e.g. Lecideion tumidae, Rhizocarpon alpicolae, and Umbilicarion<br />
cylindricae; list IV). As saxicoles Micarea species are more prevalent in sheltered ravines and<br />
woodlands. In dry underhangs in rock faces, steep banks, and below overhanging trees, up to<br />
seven species (Hst I) may be found in <strong>the</strong> ombrophobous, aerohygrophilous Micareetum<br />
sylvicolae. The species in this community grow on rock, loose or lodged stones, exposed roots,<br />
consolidated soil, and encroaching dry mats <strong>of</strong> bryophytes. Associated lichens from o<strong>the</strong>r<br />
genera may include Coniocybe furfuracea, Enterographa hutchinsae, Melaspilea subarenacea,<br />
Microcalicium arenarium, Opegrapha gyrocarpa, O. zonata, Porina chlorotica, P. lectissima,<br />
Psilolechia clavulifera, and P. lucida. This community is usually well defined, but in some<br />
situations it intergrades with o<strong>the</strong>r assemblages <strong>of</strong> shaded rocks such as <strong>the</strong> Lecideetum lucidae,<br />
Coniocybetum fururaceae, Opegraphetum horistico-gyrocarpae, and <strong>the</strong> Racodietum rupestris.<br />
A few Micarea species (list II) may be found on <strong>the</strong> upper sides <strong>of</strong> stones, boulders, and rocks<br />
where <strong>the</strong>y are subjected to direct wetting by rain, or by drips from <strong>the</strong> overlying tree canopy. In<br />
most cases <strong>the</strong>se are incidental occurrences <strong>of</strong> species more characteristic <strong>of</strong> o<strong>the</strong>r substrata.<br />
However, such situations may be <strong>the</strong> normal habitat <strong>of</strong> <strong>the</strong> little known M. curvata and M.<br />
lithinella.<br />
In nor<strong>the</strong>rn and western Britain several Micarea species occur on mossy rocks. The communities<br />
involved are <strong>of</strong>ten difficult to place, some belonging to bryophyte-dominating syntaxa,<br />
o<strong>the</strong>rs to normally epiphytic communities (e.g. Parmelietum laevigatae), or to chomophytic<br />
communities dominated by bryophytes and Cladonia species which, in turn, are <strong>of</strong>ten invested<br />
by a gelatinous algal scum.<br />
Terricolous habitats<br />
Areas <strong>of</strong> lowland heathlands where <strong>the</strong> peaty or sandy soil has been laid bare <strong>of</strong> tall vegetation<br />
by erosion, disturbance, or burning are colonised by several species <strong>of</strong> algae bryophytes and<br />
lichens. The lichens involved <strong>of</strong>ten include species such as Lecidea icmalea, L. uliginosa, L.<br />
oligotropha (rare in Britain), Baeomyces rufus, B. roseus, and some Cladonia spp. Micarea<br />
species are not usually involved in such communities, although M. melaena may at times be<br />
present, occasionally achieving local dominance. In mature Callunetum in <strong>the</strong> heathlands <strong>of</strong><br />
eastern England and Jylland (Denmark) M. nitschkeana is sometimes encountered in abundance<br />
on litter, as well as on <strong>the</strong> thin attached twigs <strong>of</strong> <strong>the</strong> Calluna. Acidic soils contaminated by<br />
heavy metals (especially lead) are <strong>of</strong>ten ra<strong>the</strong>r bare with an open vegetation, and Micarea<br />
species (especially M. lignaria) may occur in quantity growing over moribund bryophytes and<br />
plant debris. Soil, plant debris, and moribund bryophytes amongst coastal rocks are <strong>of</strong>ten<br />
colonized by M. prasina (q.v.). M. melaenida (q.v.) is found exclusively on consohdated fine<br />
grained (argillaceous) mineral soils. The terricolous species encountered in upland, montane, or<br />
arctic regime are given in list V; <strong>the</strong> reader is referred to <strong>the</strong> individual species accounts for<br />
fur<strong>the</strong>r details.<br />
Man-made substrata<br />
A detailed treatment <strong>of</strong> <strong>the</strong> lichens <strong>of</strong> man-made substrata is given by Brightman & Seaward<br />
(1977). Several Micarea species occur on worked wood (list X), but M. denigrata {q. v. ) is by far<br />
<strong>the</strong> most successful species. M. denigrata has also been found on pieces <strong>of</strong> hardboard lying in a<br />
dune slack at Tentsmuir in Fife. Old sackcloth and o<strong>the</strong>r fabrics lying on <strong>the</strong> ground in old<br />
lead-mine workings are frequently colonised by lichens, including M. lignaria. A more surpris-<br />
ing find was that <strong>of</strong> M. nitschkeana, growing with Scoliciosporum umbrinum, on a small plastic<br />
carton in a heathland on <strong>the</strong> Isle <strong>of</strong> Wight. Fur<strong>the</strong>r field studies will undoubtedly extend this list<br />
<strong>of</strong> artificial substrata. Micarea denigrata and M. prasina are rapid colonisers <strong>of</strong> newly available<br />
substrata, and have been collected respectively on dead culms <strong>of</strong> Cladium mariscus and<br />
Phragmites australis in natural habitats. Both <strong>of</strong> <strong>the</strong>se reeds are sometimes used as thatch, and a<br />
close inspection <strong>of</strong> thatched ro<strong>of</strong>s ought to reveal <strong>the</strong> presence <strong>of</strong> Micarea species.
LICHEN GENUS MICAREA IN EUROPE 93<br />
Habitats in lowland pastoral areas<br />
The lowlands <strong>of</strong> eastern England, <strong>the</strong> English south midlands and parts <strong>of</strong> eastern Scotland<br />
contain few 'natural' habitats suitable for Micarea species. In addition, <strong>the</strong>se same areas suffer,<br />
at least to some extent, from <strong>the</strong> effects <strong>of</strong> incoming pollution (especially sulphur dioxide and its<br />
derivatives) and indigenous pollution in <strong>the</strong> form <strong>of</strong> agricultural biocides and fertilisers.<br />
Never<strong>the</strong>less, <strong>the</strong> diligent collector will usually discover suitable niches where Micarea species<br />
are to be found. The most commonly encountered member <strong>of</strong> <strong>the</strong> genus in lowland agricultural<br />
and suburban districts is M. denigrata which can grow on worked wood in a wide range <strong>of</strong><br />
situations, and on old tree stumps in hedgerows. M. prasina can be found on trees and shrubs in<br />
small woodlands planted as fox coverts or for <strong>the</strong> rearing <strong>of</strong> pheasants. It may also be found on<br />
sheltered stems <strong>of</strong> shrubs in waste ground, marginal land, or gardens, where M. nitschkeana may<br />
also occur.<br />
A major refuge for wildlife (including lichens) in lowland Britain is provided by churchyards<br />
(Anon, 1973, 1978). Although records <strong>of</strong> Micarea species are ra<strong>the</strong>r few, M. denigrata can <strong>of</strong>ten<br />
be found on timber and, more rarely, on siliceous memorial stones; and M. lignaria and M.<br />
peliocarpa have been recorded on <strong>the</strong> siliceous stone-work <strong>of</strong> old walls and memorials.<br />
Distribution<br />
Britain<br />
Maps 1-28 present <strong>the</strong> distributions <strong>of</strong> all <strong>the</strong> Micarea species occurring in <strong>the</strong> <strong>British</strong> Isles, with<br />
exception <strong>of</strong> M. lithinella which is known from just one 10 km grid square (44/86). It is seen from<br />
<strong>the</strong>se maps that records for most species are concentrated in <strong>the</strong> western and/or nor<strong>the</strong>rn<br />
districts. This is partly due to <strong>the</strong> fact that suitable habitats are most numerous in <strong>the</strong>se areas,<br />
and it is probable that several species (e.g. M. bauschiana, M. lignaria, M. melaena, M.<br />
peliocarpa, and M. sylvicola) would be more evenly distributed if <strong>the</strong>re were more available<br />
habitats in <strong>the</strong> lowland areas <strong>of</strong> <strong>the</strong> south and east. It is impossible to discuss in isolation <strong>the</strong><br />
effects <strong>of</strong> climate or substrate availability on plant distributions. Substrate availability is largely<br />
dependent on topography and geology, but <strong>the</strong> physical and chemical nature <strong>of</strong> an existing<br />
substratum can be much influenced by <strong>the</strong> prevailing climate. For example, <strong>the</strong> bark <strong>of</strong> trunks <strong>of</strong><br />
mature Quercus in high rainfall districts tends to be more leached, s<strong>of</strong>t, and friable (i.e. more<br />
suitable for Micarea spp.) than in low rainfall districts. As a second example <strong>the</strong> species <strong>of</strong> <strong>the</strong><br />
Micareetum sylvicolae (see p. 92) are ombrophobous (avoid frequent direct wetting), but are<br />
also aerohygrophilous (require ± constant high humidity). In western Britain <strong>the</strong> high rainfall,<br />
high incidence <strong>of</strong> cloud cover (i.e. low duration <strong>of</strong> possible sunshine), and incoming moist<br />
air-stream from <strong>the</strong> Atlantic on <strong>the</strong> prevailing westerly winds result in high levels <strong>of</strong> relative<br />
humidity (and low levels <strong>of</strong> saturation deficit) more or less throughout <strong>the</strong> year (see Climato-<br />
logical Atlas <strong>of</strong> <strong>the</strong> <strong>British</strong> Isles, London: HMSO (1952)). Under such conditions a well<br />
developed Micareetum sylvicolae can be found on underhangs in a wide range <strong>of</strong> lowland<br />
habitats. Eastern districts generally have a lower rainfall and lower incidence <strong>of</strong> cloud cover<br />
resulting in lower levels <strong>of</strong> relative humidity, such that <strong>the</strong> Micareetum sylvicolae tends to be<br />
more confined to narrow river valleys with a long history <strong>of</strong> continuous tree cover. A western<br />
bias to <strong>the</strong> present day <strong>British</strong> distribution <strong>of</strong> many lichens is <strong>of</strong>ten due to <strong>the</strong> fact that <strong>the</strong> eastern<br />
side <strong>of</strong> <strong>the</strong> country is generally that most affected by industrial and urban development,<br />
intensive agricultural practices, and <strong>the</strong> resultant forms <strong>of</strong> air pollution (Hawksworth, Coppins<br />
& Rose, 1974; Coppins, 1976).<br />
A climatically determined western distribution, attributable to <strong>the</strong> General Western Group <strong>of</strong><br />
Coppins (1976), is shown by Micarea adnata, M. alabastrites, M. cinerea, M. lignaria var.<br />
species have a eu- Atlantic or sub-Atlantic<br />
endoleuca, M. stipitata, and M. syno<strong>the</strong>oides . These<br />
European distribution, and are confined to areas with an annual rainfall <strong>of</strong> over 800 m (mostly<br />
over 1000 mm) distributed over at least 160 wet days ['wet day' = period <strong>of</strong> 24 h in which 1 mm<br />
(or more) <strong>of</strong> rain is recorded]. The genus Micarea is poorly represented in sou<strong>the</strong>rn (Mediterranean)<br />
Europe, and it is not surprising that few <strong>British</strong> species exhibit a marked sou<strong>the</strong>rn<br />
tendency. The best example <strong>of</strong> such a species is M. pycnidiophora, which is mainly confined to
LICHEN GENUS MICAREA IN EUROPE 95<br />
The identification (and subsequent annotation and cataloguing) <strong>of</strong> a single specimen <strong>of</strong><br />
Micarea (which more <strong>of</strong>ten than not is fragmentary or in poor condition) can be very time<br />
consuming, and <strong>of</strong>ten involves lengthy microscopical examination. The unfortunate consequence<br />
<strong>of</strong> this is that I have not been able to examine a large number <strong>of</strong> potentially available<br />
specimens from several major institutional and private herbaria.<br />
Despite <strong>the</strong> above problems and shortcomings, I have tentatively attempted below to assign<br />
<strong>the</strong> species <strong>of</strong> Micarea to some general distributional types. In most cases I have paid little<br />
attention to purely literature sources <strong>of</strong> records because I have found that many such reports,<br />
even those by lichenologists whose works I hold in <strong>the</strong> highest esteem, can be unreliable in<br />
respect <strong>of</strong> current species concepts.<br />
A. Eu-Atlantic - mainly confined to oceanic areas including west Norway, <strong>the</strong> <strong>British</strong> Isles, western<br />
France, and Macaroenesia.<br />
M. alabastrites M. stipitata<br />
M. nigella M. subnigrata<br />
M. olivacea tM. subviolascens<br />
*M. prasina (with 'unknown C) M. syno<strong>the</strong>oides<br />
*M. pycnidiophora<br />
*with sou<strong>the</strong>rn tendency twith affinity to group G<br />
B. Sub-Atlantic - as above, but also present in high rainfall (annual rainfall <strong>of</strong> >800 mm) areas <strong>of</strong><br />
central Europe (Alps, Carpathians, etc.).<br />
M. adnata M. lignaria var. endoleuca<br />
M. cinerea<br />
C. Boreal - mainly confined to Fennoscandia, but absent or rare from west Norway.<br />
M. anterior M. melanobola<br />
M. contexta M. muhrii<br />
M. eximia M. osloensis<br />
M. melaeniza M. rhabdogena<br />
D. Boreal-continental - present in Fennoscandia and central Europe, but rare or absent from west<br />
Norway and western Britain.<br />
M. elachista M. lithinella<br />
M. hedlundii *M. melaenida<br />
*but known from western France<br />
E. Continental - known only from central Europe.<br />
M. curvata<br />
F. Montane (arctic-alpine) - at high altitudes in montane regions <strong>of</strong> Britain and central Europe, but<br />
sometimes at low altitudes in Fennoscandia.<br />
M. assimilata M. subleprosula<br />
M. crassipes M. turfosa<br />
M. incrassata<br />
G. Arctic - mainly confined to within <strong>the</strong> Arctic Circle, but possibly extending southwards along <strong>the</strong><br />
Atlantic coast.<br />
M. ternaria<br />
H. Widely Distributed - known from <strong>British</strong> Isles, Fennoscandia, central Europe, and <strong>of</strong>ten elsewhere.<br />
More records will probably reveal distinct phytogeographical tendencies for some <strong>of</strong> <strong>the</strong><br />
species included here.<br />
M. bauschiana M. melaena<br />
M. botryoides tM. misella<br />
M. denigrata M. myriocarpa<br />
XM. globulosella M. nitschkeana<br />
M. intrusa XM. peliocarpa<br />
M. leprosula *M. prasina<br />
M. lignaria var. lignaria<br />
M. sylvicola<br />
M. lutulata<br />
*races containing 'prasina unknowns A and B'<br />
twith possible affinity to group D<br />
twith possible affinity to group B<br />
M. tuberculata
96 BRIAN JOHN COPPINS<br />
World<br />
The present study is confined to species <strong>of</strong> Micarea that occur in Europe. However, some<br />
extra-European specimens have been examined in connection with nomenclatural matters, and<br />
o<strong>the</strong>rs have been examined incidentally. According to current information <strong>the</strong> genus is best<br />
represented in Europe, but this may, or may not, be true. Micarea is well represented in North<br />
America from where I have seen 12 species (all European taxa: M. crassipes, M. denigrata, M.<br />
globulosella, M. lignaria s. sir., M. melaena, M. misella, M. nitschkeana, M. peliocarpa, M.<br />
prasina, M. sylvicola, M. ternaria, and M. turfosa); this list will undoubtedly be much extended<br />
in <strong>the</strong> near future. From o<strong>the</strong>r regions I have seen (but not necessarily critically examined)<br />
specimens <strong>of</strong> Micarea from Japan, Borneo, Tasmania, New Zealand, South Africa, South<br />
America (Brazil), and <strong>the</strong> Antilles (Cuba). It is likely that species <strong>of</strong> Micarea are to be found in<br />
most temperate and boreal regions, as well as in many tropical regions (especially mountainous<br />
areas). The numerous collections that I have received from New Zealand include several<br />
undescribed taxa but <strong>the</strong>y also include at least one European species, Micarea peliocarpa. Three<br />
additional European species known from <strong>the</strong> sou<strong>the</strong>rn hemisphere are M. incrassata (Ker-<br />
guelen), M. lignaria (Brazil), and M. misella (Brazil).<br />
Micarea Fr.<br />
The genus Micarea<br />
Syst. orb.: 256 (1825). - Micarea Fr. emend Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18(3): 27<br />
(1892). Lectotype: Micarea prasina Fr. (see note below).<br />
Helocarpon Th. Fr., Lich. arctoi: 178 (1860); Nova Acta R. Soc. Sclent. Upsal. Ill, 3: 278 (1861). Type<br />
species: Helocarpon crassipes Th. Fr. [= Micarea crassipes (Th. Fr.) Coppins].<br />
Stereocauliscum Nyl. in Flora, Jena 48: 211 (1865). Type species: Stereocauliscum gomphillaceum Nyl.<br />
[= Micarea lignaria (Ach.) Hedl.]<br />
Micarea sect. Bryophagae Poelt & Dobbeler in Bot. Jb. 96: 337 (1975). Type species: Micarea polytrichi<br />
Poelt & Dobbeler [= Micarea prasina Fr.].<br />
Note. The name Micarea was first validly published in December 1825 by Fries {op. cit.), although it was<br />
twice mentioned by <strong>the</strong> same author earlier in <strong>the</strong> same year. In Sched. crit. lich. siiec. part 3, fasc. 4, page<br />
21 (pre 7 May) it appeared in a note under <strong>the</strong> entry for Biatorafuliginea. However, it was not accepted in<br />
that work, and must <strong>the</strong>refore be considered invalid according to Art. 34.1. The second appearance was in<br />
Stirp. agri. femsion. page 37 (June 1825) in <strong>the</strong> form <strong>of</strong> <strong>the</strong> combinations 'Micarea fuliginea and 'M. nigra'.<br />
Nei<strong>the</strong>r <strong>of</strong> <strong>the</strong>se was accompanied by a description and <strong>the</strong> generic name was again invalidly published<br />
(Arts 32, 34.1(e)).<br />
When validly published, Micarea was introduced with four species: M. prasina, M. coccinea, M.<br />
fuliginea, and M. nigra. Korber (1855) emended <strong>the</strong> genus to include only M. prasina and is <strong>the</strong>reby<br />
considered to have lectotypified <strong>the</strong> genus on that species. See 'Excluded taxa' for notes on <strong>the</strong> remaining<br />
three original names.<br />
Thallus crustose or immersed in <strong>the</strong> substratum, effuse and <strong>of</strong>ten wide-spreading, never<br />
bordered by delimiting hypothalline lines. Superficial thallus consisting <strong>of</strong> ± spherical granules<br />
(goniocysts) up to c. 60 ±m diam; or convex to subglobose areolae which in some species may<br />
dissolve into soredia: or more rarely present as a thin ± smooth to rmose, or scurfy-granular<br />
crust. Thallus in section ecorticate (or weakly corticate in a few species), but sometimes (when<br />
areolate) covered by a hyaline amorphous layer, and outermost hyphae sometimes pigmented.<br />
Phycobiont 'grass-green'; cells usually thin-walled and c. 4-7 pm ('micareoid'), or more rarely<br />
larger ('non-micareoid' phycobiont types); a few species also have cephalodia containing Nostoc<br />
ox Stigonema.<br />
Apo<strong>the</strong>cia small (mostly < 1 mm diam) , whitish or variously coloured (mostly grey , dull brown<br />
or blackish), epruinose, usually immarginate or ± so, adnate, sessile or rarely stipitate, convex<br />
to ± globose and <strong>of</strong>ten becoming tuberculate. Hymenium with amyloid gel-matrix. Asci clavate<br />
or cylindrical-clavate, <strong>of</strong> Lecanora-type, 8-spored. Spores hyaline, smooth-walled, variously<br />
shaped (ellipsoid, ovoid, fusiform, or acicular), usually less than 6 pm wide, simple to<br />
multiseptate but never muriform. Paraphyses few to numerous, septate, mostly branched<br />
(especially above), <strong>of</strong>ten anastomosing, mostly in range <strong>of</strong> 0-7-T7 pm wide at mid-hymenium;
LICHEN GENUS MICAREA IN EUROPE 97<br />
apices never regularly clavate or capitate and never with a dark brown apical 'cap', but<br />
sometimes irregularly incrassate and sometimes with thickened pigmented walls in about upper<br />
5-15 ixm; in addition, some species have a few stout paraphyses (which are c. 2-3 /xm wide, and,<br />
in some cases, pigmented throughout <strong>the</strong>ir length) occurring as scattered individuals or small<br />
fascicles. Hypo<strong>the</strong>cium (including subhymenium) variously pigmented, <strong>of</strong> interwoven hyphae<br />
that become outwardly orientated towards <strong>the</strong> hymenium, mixed with wider, short-celled<br />
ascogenous hyphae. Excipulum <strong>of</strong>ten absent or indistinct, if discernible <strong>the</strong>n non-amyloid (or ±<br />
so) and composed <strong>of</strong> radiating branched and anastomosing paraphysis-like hyphae that become<br />
distinct and ± separate in K.<br />
Pycnidia <strong>of</strong>ten present, very varied in form from immersed to sessile or stalked, <strong>the</strong> stalks<br />
(pycnidiophores) sometimes branched. Pycnidial walls hyaline or pigmented. Conidiogenous<br />
cells ampulliform to cylindrical (sometimes with swollen base), phialidic, sometimes with 1-3<br />
proliferations. Conidia hyaline, smooth-walled, <strong>of</strong> three basic types: (i) microconidia - ±<br />
cylindrical, aseptate, eguttulate, in range 3-5-9x0-5-l fxm, borne in immersed to sessile<br />
pycnidia usually
98 BRIAN JOHN COPPINS<br />
species in at least one case. Micarea intrusa {q.v.) possibly provides a link between <strong>the</strong> two<br />
genera, and its inclusion in Micarea, ra<strong>the</strong>r than Scoliciosporum, is mainly on <strong>the</strong> grounds <strong>of</strong><br />
spore-type. The differences between <strong>the</strong> two genera are admittedly unclear and ill-defined at <strong>the</strong><br />
present time, and will certainly require re-appraisal as our knowledge <strong>of</strong> morphological,<br />
ontogenetic, and phylogenetic aspects in <strong>the</strong> Lecideaceae increases.<br />
A genus which may be closely related to both Micarea and Scoliciosporum is Strangospora<br />
Massal.; it shares with <strong>the</strong>m unsophisticated apo<strong>the</strong>cial and thallus structures, and similar<br />
ecological requirements. However, Strangospora differs in having polysporous asci with aseptate,<br />
globose spores, although <strong>the</strong> asci do appear to be <strong>of</strong> <strong>the</strong> Lecanora-typQ. The monotypic<br />
genus Steinia Korber is represented by S. geophana (Nyl.) Stein, whose small, black, immargin-<br />
ate apo<strong>the</strong>cia occurring on shaded substrate (soil, rotten wood, rocks, stones, etc.) are easily<br />
confused in <strong>the</strong> field with those <strong>of</strong> some Micarea species. However, it is easily distinguished in<br />
microscopical preparations by its 16-spored asci with globose spores, and simple, slender<br />
paraphyses. In addition, its asci are not <strong>of</strong> <strong>the</strong> Lecanora-typQ, although <strong>the</strong>ir apices are each<br />
provided with a broad, amyloid plug.<br />
Micarea appears to have some affinities to Psilolechia Massal., and <strong>the</strong> two genera are<br />
compared in <strong>the</strong> discussion <strong>of</strong> Psilolechia clavulifera (p. 376). Micarea may also be close to <strong>the</strong><br />
predominantly foliicolous Byssoloma Trevisan, but species <strong>of</strong> this latter genus have an excipulum<br />
<strong>of</strong> loosely woven, hyaline, pachydermatous hyphae which spread laterally to form a<br />
conspicuous white border to <strong>the</strong> apo<strong>the</strong>cium. However, some specimens <strong>of</strong> B. subdiscordans<br />
(Nyl.) P. James, when growing on bark, mosses, or rocks, do not have conspicuous whitebordered<br />
apo<strong>the</strong>cia, although <strong>the</strong> pachydermatous hyphae are clearly seen in sections <strong>of</strong> <strong>the</strong><br />
apo<strong>the</strong>cia.<br />
Lignicolous members <strong>of</strong> <strong>the</strong> 'Lecanora' symmicta group are occasionally confused with<br />
Micarea species on account <strong>of</strong> <strong>the</strong>ir general appearance and excipulum structure. Their<br />
excipular hyphae are radiating, richly branched and anastomosed and lax in K, although <strong>the</strong>y<br />
contrast markedly with <strong>the</strong> simple or sparingly branched paraphyses and do not give <strong>the</strong><br />
impression <strong>of</strong> being 'paraphysis-like'. This group fur<strong>the</strong>r differs from Micarea in having dense<br />
epi<strong>the</strong>cial granules which dissolve in K and a generally different suite <strong>of</strong> Uchen substances (which<br />
includes usnic acid and zeorin). Ano<strong>the</strong>r seemingly distinctive group <strong>of</strong> species (but not yet<br />
afforded generic status) that bears some close resemblances to Micarea is that containing such<br />
species as Catillaria contristans, Lecidea limosa, and L. stenotera; for fur<strong>the</strong>r discussion see<br />
account oi Micarea assimilata (p. 115).<br />
The apo<strong>the</strong>cia <strong>of</strong> Vezdaea species are superficially similar to those <strong>of</strong> some species oi Micarea,<br />
but differ in that <strong>the</strong>ir tissues are not bound by a gelatinous matrix and <strong>the</strong>ir asci have uniformly<br />
amyloid walls and do not belong to <strong>the</strong> Lecanora-type; see Poelt & Dobbeler (1975).<br />
There has been some confusion between Micarea and genera such as Arthonia Ach., and<br />
Chrysothrix Mont., and also <strong>the</strong> Trapelia-\ike 'Lecidea' granulosa group. These all differ from<br />
Micarea in not having Lecanora-typc asci (see 'Excluded taxa').<br />
Suprageneric considerations<br />
The formulation <strong>of</strong> a ± stable hierarchical classification <strong>of</strong> fungal (including lichen) taxa above<br />
<strong>the</strong> rank <strong>of</strong> genus is unUkely to be achieved for some decades. A suprageneric classification for<br />
<strong>the</strong> lichenized fungi has been attempted in recent years by, for example , Henssen & Jahns (1973)<br />
and Poelt (19746). These schemes are extremely useful bases for fur<strong>the</strong>r study but, as admitted<br />
by <strong>the</strong>ir authors, contain very many uncertainties and points <strong>of</strong> conjecture, and should be<br />
regarded as being <strong>of</strong> a very provisional nature.<br />
Micarea has Lecanora-type asci and clearly belongs in <strong>the</strong> order Lecanorales and suborder<br />
Lecanorineae as defined by Poelt (19746). My belief is that Micarea should be placed in <strong>the</strong><br />
Lecideaceae Chev. s. sir. (i.e. confined to species with Lecanora-type asci), with <strong>the</strong> provision<br />
that <strong>the</strong> real differences (if any) with that family and <strong>the</strong> Lecanoraceae Fee need to be explored.<br />
For <strong>the</strong> time being (at least) I cannot accept <strong>the</strong> 'Micareaceae Vezda ad int.' (Poelt, 19746: 627;<br />
Eriksson, 1981), although it may be possible to formally define this family name in <strong>the</strong> future as<br />
knowledge and taxonomic concepts in <strong>the</strong> Lecanorales increases and advances.
Infrageneric considerations<br />
LICHEN GENUS MICAREA IN EUROPE 99<br />
I have given much thought to <strong>the</strong> possibihty <strong>of</strong> subdividing <strong>the</strong> genus Micarea into subgenera or<br />
sections. The genus includes several small groups <strong>of</strong> closely related species, but <strong>the</strong> affinities <strong>of</strong><br />
many individual species are difficult to ascertain. I believe that a formal infrageneric classifica-<br />
tion (above <strong>the</strong> rank <strong>of</strong> species) based on current information would be unhkely to withstand <strong>the</strong><br />
tests <strong>of</strong> time and serve httle useful purpose. However, that is not to say that such a classification<br />
will never be possible. The present study is mainly confined to <strong>the</strong> 45 European species <strong>of</strong><br />
Micarea, but <strong>the</strong> genus is well represented in o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong> world by some <strong>of</strong> <strong>the</strong> same, plus<br />
numerous additional species. With such considerations in mind I estimate that <strong>the</strong> genus, as<br />
currently circumscribed, contains about 100 species world-wide. A better understanding <strong>of</strong><br />
extra-European species may increase <strong>the</strong> feasibility <strong>of</strong> subdividing Micarea.<br />
Some examples <strong>of</strong> small groups <strong>of</strong> apparently closely related species are given below. Within<br />
each group all <strong>the</strong> species have basically similar apo<strong>the</strong>cial construction. For example. Group C<br />
all have medium to large sized apo<strong>the</strong>cia, a ± well-developed excipulum, and numerous<br />
paraphyses which are <strong>of</strong> medium thickness and are richly branched (especially in <strong>the</strong> upper<br />
hymenium). Some supplementary common features and o<strong>the</strong>r notes are provided for each<br />
group - but for more detailed discussions see <strong>the</strong> taxonomic accounts <strong>of</strong> <strong>the</strong> relevant species.<br />
Phycobiont is 'micareoid' unless o<strong>the</strong>rwise stated. See under 'Chemistry' for explanation <strong>of</strong><br />
pigments. Abbreviations: AL, hyaUne amorphous covering layer; Exc, excipulum; Hym.,<br />
hymenium; Hyp., hypo<strong>the</strong>cium; Th., thallus; Pyc, pycnidia.<br />
A. M. lignaria - M. ternaria<br />
Th. areolate-type with AL. Th., Hym. and Pyc. with pigment A; Hyp. with dilute pigment A plus<br />
dilute dull brown pigment. Spores ± fusiform, 3- or more septate. Mesoconidial states very similar;<br />
M. lignaria also has micro- and macroconidial states. Chemistry: argopsin or xanthones (M.<br />
lignaria) or no substances {M. ternaria).<br />
B. M. leprosula-M. subleprosula<br />
Th. sorediate-type. Th. and Hym. with pigment A; Hyp. <strong>of</strong>ten with dilute dull brown pigment.<br />
Spores ± fusiform, 3- or more septate. Anamorphs unknown. Chemistry: argopsin -I- gyrophoric<br />
acid (M. leprosula) or alectorialic acid -I- accessory substances (M. subleprosula). Many affinities<br />
with Groups A and C.<br />
C. M. peliocarpa - M. alabastrites - M. cinerea<br />
Th. areolate-type with AL. Hym. , Th. and Pyc. <strong>of</strong> M. peliocarpa and M. cinerea with pigment A;<br />
all parts <strong>of</strong> M. alabastrites devoid <strong>of</strong> pigment. Spores ± fusiform, 3- or more septate. All with<br />
microconidia and curved or filiform macroconidia; mesoconidia unknown. Chemistry: all with<br />
gyrophoric acid. With Atlantic or sub-Atlantic distributions. Some affinities with Groups A, B, and<br />
D.<br />
D. M. denigrata - M. nitschkeana - M. globulosella<br />
Th. areolate-type but with no AL. Hym., Th. and Pyc. with pigment D; Hyp. hyaline. All with<br />
similar micro- and mesoconidial states; M. denigrata and M. nitschkeana also with ± identical<br />
macroconidial states. Chemistry: all with gyrophoric acid. Mainly corticolous or lignicolous. These<br />
three species appear to show a good example <strong>of</strong> an evolutionary progression: <strong>the</strong>y are morphologi-<br />
cally and chemically ± identical except for <strong>the</strong> length and septation <strong>of</strong> spores.<br />
E. M. pycnidiophora - M. stipitata<br />
Th. areolate-type, but with no AL. Without pigments. Spores ± acicular, 3-7-septate. Pyc.<br />
stalked, with mesoconidia; micro- and macroconidia unknown. Chemistry: gyrophoric acid (Af.<br />
pycnidiophora) or no substances (M. stipitata). Corticolous and with eu-Atlantic distributions.<br />
F. M. elachista - M. rhabdogena<br />
Th. areolate-type with cortex and AL (M. elachista) or endoxylic (M. rhabdogena). Upper Hym.<br />
with pigment E; Pyc. with pigment D; Hyp. hyaline. Macroconidia unknown. Chemistry: no<br />
substances. Mainly lignicolous with boreal or boreal-continental distribution.<br />
G. M. botryoides-M. melaeniza<br />
Th. weakly areolate (with no AL) or with goniocysts (Af. botryoides) , or endoxylic {M. melaeniza).<br />
Hym. and Pyc. walls with pigment A; Hyp. and Pyc. stalks with pigment F. Paraphyses dimorphic.<br />
Pyc. stalked with mesoconidia; micro- and macroconidia unknown. Chemistry: no substances. This
100 BRIAN JOHN COPPINS<br />
group could possibly be extended to include M. anterior, and <strong>the</strong>re are probably some affinities to<br />
species <strong>of</strong> Groups H and I.<br />
H. M. contexta - M. eximia - M. nigella - M. olivacea<br />
Th. endoxylic, or forming a thin crust, or weakly areolate. Pigments A and (or) B in various<br />
locations. Spores simple or 1-septate. Paraphyses dimorphic. Pyc. black, immersed to sessile, or<br />
stalked (A/, nigella). Macroconidia unknown. Chemistry: no substances. Mainly lignicolous and<br />
confined (?) to north-west Europe. Probably close to Group G. M. melaena may belong here but<br />
has: ra<strong>the</strong>r large apo<strong>the</strong>cia, pigment C (usually present in hypo<strong>the</strong>cium), an <strong>of</strong>ten well-developed,<br />
areolate thallus containing gyrophoric acid, and an ability to produce macroconidia; it may provide<br />
a link between Group H and Groups A-D.<br />
I. M. bauschiana - M. sylvicola - M. tuberculata - M. lutulata<br />
Th. weakly areolate, ± smooth or scurfy. Phycobiont non-micareoid. Pigmentation variable,<br />
involving pigments A, B, C, and F; pigment D never present. Spores small, simple, or p.p.<br />
1-septate (M. tuberculata). Paraphyses dimorphic. Pyc. ± immersed. Macroconidia unknown.<br />
Chemistry: no substances. Mainly found in dry underhangs in <strong>the</strong> Micareetum sylvicolae. This<br />
group is almost worthy <strong>of</strong> subgeneric status but it shows some affinities to Group G, and o<strong>the</strong>r<br />
species, such as M. lithinella.<br />
J. M. assimilata - M. incrassata - M. subviolascens - M. melaenida - M. crassipes<br />
Th. areolate-type, sometimes with AL. Pigmentation variable, involving pigments A, B, C, D, and<br />
E; Hyp. always dark-coloured. Spores ra<strong>the</strong>r large, mostly ellipsoid or oblong-elhpsoid and simple<br />
or 1-septate. Paraphyses ra<strong>the</strong>r stout and sparingly branched. Cephalodia found in first three<br />
species. Pyc. immersed. Macroconidia unknown. Chemistry: no substances. Mainly terricolous or<br />
muscicolous (never corticolous or lignicolous), and with a ± arctic or arctic-alpine distribution<br />
(except M. melaenida). M. crassipes is ra<strong>the</strong>r anomalous here because <strong>of</strong> its very well-developed<br />
excipulum and turbinate or stipitate apo<strong>the</strong>cia.<br />
K. M. prasina - M. hedlundii - 'Lecidea' levicula [from Cuba]<br />
Th. goniocyst-type. Hym., Pyc. and Th. <strong>of</strong>ten with pigment D; Hyp. hyaUne; one species (M<br />
hedlundii) with pigment H. Spores mostly simple or 1-septate. Pyc. immersed to sessile, or stalked<br />
and tomentose {M. hedlundii). Micro- and (or) mesoconidia produced; macroconidia unknown.<br />
Chemistry: variable, including 'prasina unknowns' and gyrophoric acid. M. misella and M.<br />
melanobola have some affinities to this group, but also with Group D. M. syno<strong>the</strong>oides may belong<br />
near here.<br />
Keys to species<br />
Guide to keys and identifications<br />
Two keys are provided, <strong>the</strong> first to specimens with apo<strong>the</strong>cia, <strong>the</strong> second to specimens without<br />
apo<strong>the</strong>cia (although <strong>of</strong>ten with pycnidia). The keys are for European species, but all those<br />
known to me from cool-temperate, boreal, and arctic regions <strong>of</strong> North America are, by chance,<br />
included.<br />
The nature <strong>of</strong> pigmentation and colour reactions observed in anatomical sections is important<br />
for <strong>the</strong> identification <strong>of</strong> Micarea species. Many <strong>of</strong> <strong>the</strong> subtle yet significant colour hues and<br />
reactions are obscured by <strong>the</strong> yellowish illumination given by most bulbs, and it is recommended<br />
that a microscope be fitted with a 'daylight' bulb or a blue filter. The preceding sections on<br />
'Morphology' and 'Chemistry' should be read before attempting to use <strong>the</strong> keys.<br />
Particular care should be taken in determining C reactions with apo<strong>the</strong>cial sections. The straw<br />
to dull olivaceous, K+ violet pigment found, for example, in M. prasina, M. syno<strong>the</strong>oides, M.<br />
denigrata, and M. nitschkeana, also reacts C+ violet (persistent); this reaction tends to<br />
mask <strong>the</strong> C+ orange-red (quickly fading) reaction due to gyrophoric acid and usually obtainable<br />
in <strong>the</strong> last two, aforementioned species. However, <strong>the</strong> C+ violet reaction is mostly confined to<br />
<strong>the</strong> upper part <strong>of</strong> <strong>the</strong> hymenium, whereas <strong>the</strong> C+ orange-red reaction occurs in all parts <strong>of</strong> <strong>the</strong><br />
apo<strong>the</strong>cium. To carry out <strong>the</strong> C test cut a hand section <strong>of</strong> an apo<strong>the</strong>cium and mount in a drop <strong>of</strong><br />
water near <strong>the</strong> edge <strong>of</strong> <strong>the</strong> cover-slip; lay a piece <strong>of</strong> tissue-paper over <strong>the</strong> cover slip to take up any<br />
excess water; place <strong>the</strong> slide preparation on <strong>the</strong> microscope stage and focus at about x 100 to<br />
x200 and note any pigmentation; <strong>the</strong>n, apply a drop <strong>of</strong> C by <strong>the</strong> edge <strong>of</strong> <strong>the</strong> cover-slip nearest
LICHEN GENUS MICAREA IN EUROPE 101<br />
<strong>the</strong> section. A positive reaction due to gyrophoric acid is seen as an orange-red front moving<br />
across <strong>the</strong> section, with <strong>the</strong> colour quickly fading in its wake. If <strong>the</strong> concentration <strong>of</strong> gyrophoric<br />
acid is very high than <strong>the</strong> reaction may be more immediate and intense, but, never<strong>the</strong>less, <strong>the</strong><br />
colour quickly fades away within a few seconds. A similar procedure should be followed when<br />
testing for o<strong>the</strong>r colour changes.<br />
The identification <strong>of</strong> Micarea species usually requires accurate measurement <strong>of</strong> <strong>the</strong> dimen-<br />
sions <strong>of</strong> spores, conidia, and paraphyses, such that a microscope with a good resolution at<br />
X 1000, coupled with a carefully calibrated measuring eyepiece, is invaluable, and in many cases<br />
essential.<br />
Spores should be observed and measured in 10% KOH; in most cases this solution will render<br />
any septa visible. If, however, <strong>the</strong> spore contents are difficult to clear, septa can be discerned by<br />
preparing an apo<strong>the</strong>cial section or squash in LCB and heating (just to boiHng) over a spirit lamp;<br />
any septa should <strong>the</strong>n be clearly seen, especially under oil-immersion (xlOOO).<br />
Conidia should always be examined under oil-immersion (at least xlOOO). Preparations can<br />
be made in 10% KOH, but better resolution is obtained using ammoniacal erythrosin. In water,<br />
small conidia are prone to 'Brownian movement', making alignment with <strong>the</strong> measuring<br />
graticule a frustrating task! If conidia and conidiogenous cells are to be examined in a squash<br />
preparation it is <strong>of</strong>ten helpful to soak an intact pycnidium in a small drop <strong>of</strong> 10% KOH on a shde<br />
for about a minute; excess KOH is <strong>the</strong>n soaked up with <strong>the</strong> edge <strong>of</strong> a piece <strong>of</strong> tissue-paper or<br />
filter-paper and <strong>the</strong> squash prepared under a cover-slip in ammoniacal erythrosin. The KOH<br />
s<strong>of</strong>tens <strong>the</strong> pycnidial wall and does not alter <strong>the</strong> effectiveness <strong>of</strong> <strong>the</strong> erythrosin.<br />
Fine measurements, such as <strong>the</strong> width <strong>of</strong> spores, conidia, hyphae, and paraphyses should not<br />
be made in LCB, this solution <strong>of</strong>ten causing much shrinkage; with hyphae and paraphyses <strong>the</strong><br />
cytoplasm (lumina) is intensely stained, but <strong>the</strong> delimitation <strong>of</strong> <strong>the</strong>ir outer walls is <strong>of</strong>ten difficult<br />
to ascertain.<br />
Key to European species<br />
la Hymenium, atleast in upper part, dull greenish or brownish in water, K-l- violet [pigment D].. 2<br />
lb Hymenium variously coloured or hyaline, not turning to violet in K (if purple in K, <strong>the</strong>n<br />
pigment already purplish in water [pigment C] 12<br />
2a (la) Hypo<strong>the</strong>cium dark purple-brown, K+ purple intensifying or K+ green in upper part.<br />
Saxicolous 40. M. subviolascens (p. 185)<br />
2b Hypo<strong>the</strong>cium hyaline or pale . Usually corticolous or lignicolous 3<br />
3a (2b) Spores mostly 3 (or more)-septate , or over 15 /u,m long 4<br />
3b Spores mostly simple or 1-septate (2- or 3-septate spores if present very rare) , mostly less than<br />
15 /am long<br />
4a (3a) Apo<strong>the</strong>cia sections and/or thallus C + orange-red (gyrophoric acid) 5<br />
4b Apo<strong>the</strong>cia sections and thallus C- [note: <strong>the</strong> K+ violet pigment also reacts C+ violet],<br />
gyrophoric acid absent<br />
5a (4a) Spores 0-l(-2)-septate, few exceeding 16 /xm in length. Usually lignicolous. Rare forms<br />
<strong>of</strong> this very variable species 11. M. denigrata (p. 127)<br />
5b Spores mostly 3 (or more)-septate, or many exceeding 16 /am in length. Usually on bark or<br />
twigs<br />
6<br />
6a (5b) Spores (l-)3-septate, fusiform, <strong>of</strong>ten curved, 10-17(-19)x2-5-3(-3-5) /am. Microconidia<br />
(4-7-)5-5-7-5x0-8 /am; mesoconidia (3-)3-5-5(-5-7)x 1-1-5 /am; macroconidia some-<br />
times present, curved 30. M. nitschkeana (p. 165)<br />
6b Spores 0-3(-6)-septate, acicular, 13-26xl-5-2-5(-3) /am. Microconidia 3-8-5 xO-8-1 /am;<br />
mesoconidia 3-6-5-3 x 1-1-4 /am; macroconidia unknown 14. M. globulosella (p. 134)<br />
8<br />
7
102 BRIAN JOHN COPPINS<br />
7a (4b) Thallus dull grey-green (never whitish) to dark olivaceous or blackish, <strong>of</strong> discrete or<br />
coalescing granular-areolae c. 20-70 /xm diam, ± gelatinous when wet. Phycobiont<br />
micareoid, cells 4-8 /im diam. Apo<strong>the</strong>cia grey- or brown-black, epruinose. Spores<br />
l-5(-ll)-septate, 14-35(^3) xl-8-2-5(-3) fim. Microconidia and/or mesoconidia <strong>of</strong>ten<br />
present, both exceeding 3-7 fxtn in length. An oceanic species on acid bark (sometimes<br />
over bryophytes)<br />
42. M. syno<strong>the</strong>oides (p. 188)<br />
7b Thallus whitish, tinged grey or green, ± endophloeodal or thin and smooth to ± verrucose,<br />
<strong>of</strong>ten rimose; never gelatinous. Phycobiont not micareoid, cells 8-16 ixm diam. Apo<strong>the</strong>cia<br />
grey to black, <strong>of</strong>ten with whitish pruina. Spores (l-)3-7-septate, 17-26x1 •5-2-5 )u,m,<br />
Excipulum hyphae coherent in K. Conidia <strong>of</strong> one type only, 2-8-3-5xl-l-4 /xm. Widespread<br />
species on ra<strong>the</strong>r basic bark , rarely on lignum Bacidia beckhausii<br />
8a (3b, 31a) Thallus <strong>of</strong> pale to dark green goniocysts c. 12-40(-60) fxm diam, <strong>of</strong>ten ± gelatinous<br />
when wet, C-, gyrophoric acid absent or detectable in trace amounts (?contaminant) by<br />
t.l.c<br />
8b Thallus endoxylic or <strong>of</strong> whitish or grey granular-areolae c. 60-200 fim diam, not appearing<br />
gelatinous when wet, CorC-l- orange-red , gyrophoric acid <strong>of</strong>ten present 11<br />
9a (8a) Goniocysts K- or hyphae violaceous in K, pigment never oily. Pycnidia immersed or<br />
sessile, not tomentose 10<br />
9b Goniocysts containing purple oily substance in K. Pycnidia numerous, distinctly stalked,<br />
brown with white tomentum; containing mesoconidia (4-)4-5-5-5(-6)xl-3-l-7 /xm.<br />
Apo<strong>the</strong>cia usually few or absent, brown, soon tuberculate. On s<strong>of</strong>t lignum <strong>of</strong> conifers,<br />
rare<br />
15. M. hedlundii (p. 135)<br />
10a (9b) Apo<strong>the</strong>cia dark grey to black, small, 0-1-0-24 mm diam. Paraphyses with dark apical<br />
walls giving a distinct dark green, K+ violet epi<strong>the</strong>cium. Spores 0-1-septate, 7-9-7x<br />
2-5-3-3 /xm. Microconidia 4-5-5 xO-7-0-8 /xm; mesoconidia 3-3-4-5x1-1 -3 /xm.<br />
10b<br />
Thallus indistinct, <strong>of</strong> scattered olivaceous goniocysts. Rare, (?) confined to Picea bark in S.<br />
Finland<br />
25. M. melanobola (p. 156)<br />
Apo<strong>the</strong>cia white to grey, sometimes blackish, 0-1-0-4 mm diam. Without sharply delimited<br />
epi<strong>the</strong>cium, <strong>the</strong> olivaceous, K-l- violet pigment mostly confined to <strong>the</strong> gel-matrix and<br />
± diffuse although <strong>of</strong>ten more concentrated towards upper part <strong>of</strong> hymenium.<br />
Spores variable, 0-l(-3)-septate, 8-14(-17)x2-3-4(-5) /xm. Microconidia (5-)5-5-8x<br />
0-7-1 /xm; mesoconidia 3-5-4-7x1-4-1-8 /xm. Thallus <strong>of</strong> thinly<br />
goniocysts, sometimes forming a thick crust. Common and<br />
scattered to coherent<br />
widespread on bark,<br />
lignum and turf (especially coastal), rare on rock; morphologically and chemically very<br />
variable<br />
34. M. prasina (p. 173)<br />
11a (8b) Apo<strong>the</strong>cia grey to black, sometimes pallid or brown (shade forms), 0-1-0-5 mm diam.<br />
Thallus <strong>of</strong> distinct whitish areolae, or dark and scurfy (parasitized), occasionally endoxylic.<br />
Spores mostly 1-septate, <strong>of</strong>ten ± curved, (7-)9-16(-18)x2-3-3(-3-5) /xm. Paraphyses<br />
numerous, c. 1-1-5 /xm wide. Pycnidia immersed or emergent, never distinctly stalked.<br />
Microconidia (4-5-)5-7-5 x 0-7-0-8 /xm; mesoconidia 2-8-4-5(-5) x 1 -2-1-8 /im; macroconi-<br />
dia curved, 12-24x1 /xm. Apo<strong>the</strong>cia and thallus sections usually C-l- orange-red (gyrophoric<br />
acid)<br />
ll.M. denigrata(p. 127)<br />
lib Apo<strong>the</strong>cia black (very rarely paler), smaller, 0-1-0-3 mm diam. Thallus usually endoxylic,<br />
rarely forming whitish areolae. Spores mostly simple, 1-septate spores rarely numerous,<br />
never curved, 6-5-9-5x2-3(3-7) /xm. Paraphyses scanty, thin, 0-5-0-8 /xm wide. EndoxyUc<br />
forms usually with black, stalked pycnidia containing mesoconidia, 3-5-5x1-1-5 /xm;<br />
microconidia 3-8-6x0-7-0-8 /xm, borne in immersed pycnidia; macroconidia unknown.<br />
Apo<strong>the</strong>cia sections usually C-, very rarely C-l- orange-red; thallus when superficial and<br />
well developed C+ orange-red (gyrophoric acid) 26. M. misella (p. 158)<br />
12a (lb) Spores acicular or sigmoid-curved (mostly over 18 /xm long and under 3-5 /xm wide at<br />
maturity)<br />
13<br />
12b Spores not acicular or sigmoid-curved 15<br />
9
LICHEN GENUS MICAREA IN EUROPE 103<br />
13a (12a) Apo<strong>the</strong>cia whitish, without pigmentation, epruinose, C+ red or C-. Phycobiont<br />
micareoid , cells 4-7 /xm diam . Pycnidia always abundant, sessile or stalked, whitish 14<br />
13b Apo<strong>the</strong>cia dark coloured (with brownish or greenish pigment in upper hymenium); if<br />
whitish <strong>the</strong>n pruinose (epi<strong>the</strong>cium finely granular) or not. Phycobiont not micareoid,<br />
many cells over 8 ^tm diam. Pycnidia (?) unknown; if present, <strong>the</strong>n never conspicuous or<br />
stalked Scoliciosporum species<br />
[see Poelt & Vezda (1981)]<br />
14a (13a) Apo<strong>the</strong>cia and pycnidia C+ red (gyrophoric acid). Pycnidial structures mostly<br />
unbranched, 0- 1-0-3 mm tall; containing mesoconidia 4-6x1-1-5 fim<br />
[if spores fusiform and pycnidia immersed, see 23]<br />
35. M. pycnidiophora (p. 179)<br />
14b Apo<strong>the</strong>cia and pycnidia C- . Pycnidial structures <strong>of</strong>ten branched, 0-3-0-8 mm tall; containing<br />
mesoconidia 6-8x1-1-8 )u,m 37. M. stipitata (p. 182)<br />
15a (12b) Mature spores 3- or more septate, mostly over 15 fim long 16<br />
15b Mature spores simple or 1-septate 28<br />
16a (15a) Upper hymenium with distinct epi<strong>the</strong>cial layer <strong>of</strong> dense fuscous-brown pigment (without<br />
green or purple tinge), K+ dissolving into a brown solution. Spors 0-l(— 3)-septate,<br />
(9-)ll-15(-20)x2-3-5/Am. Hypo<strong>the</strong>ciumpale. Usually lignicolous 12. M. elachista (p. 131)<br />
16b Upper hymenium o<strong>the</strong>rwise, if brownish <strong>the</strong>n pigment not dissolving into a brown solution.<br />
Hypo<strong>the</strong>cium hyaline , pale or dark. Substrata various 17<br />
17a (16b) Upper hymenium purple, K-l- intensifying. Hypo<strong>the</strong>cium dark purple-brown. Rare<br />
form <strong>of</strong> 22. M. melaena (p. 150)<br />
17b Upper hymenium hyaline , greenish (sometimes tinged brown) or aeruginose 18<br />
18a (17b) Hypo<strong>the</strong>cium dark in thin section 19<br />
18b Hypo<strong>the</strong>cium hyaline or with dilute greenish or brownish tinge 20<br />
19a (18a) Hypo<strong>the</strong>cium mottled reddish brown, K-, HNO3- or + orange-brown. Upper<br />
hymenium greenish, sometimes brownish in part. Hyaline paraphyses 1-1-5 /nm wide.<br />
Thallus always C- (gyrophoric acid absent) ;terricolous 45. M. turfosa (p. 194)<br />
19b Hypo<strong>the</strong>cium dark purplish brown, K-l- green or K-l- purple intensifying (<strong>of</strong>ten a mixture <strong>of</strong><br />
both reactions, HNO3-I- purple-red. Hyaline paraphyses 0-8-1 /urn wide. Thallus when well<br />
developed C-f- red (gyrophoric acid) 22. M. melaena (p. 150)<br />
20a (18b) Apo<strong>the</strong>cia whitish, without pigmentation, C-l- red (gyrophoric acid). Spores mostly<br />
20b<br />
3-septate<br />
Apo<strong>the</strong>cia pale grey to black, <strong>of</strong>ten with a greenish tinge; if whitish <strong>the</strong>n spores mostly<br />
7-septate (M. cinerea). Upper hymenium greenish to aeruginose. Apo<strong>the</strong>cia sections C-,<br />
or C-l- orange-red (gyrophoric acid). Spores 3- or more septate<br />
21a (20a) Apo<strong>the</strong>cia usually numerous and crowded. Spores 3(-5)-septate, (16-)17-26(-28)x<br />
4-5(-6) ^tm. On bark and lignum, <strong>of</strong>ten over bryophytes, in oceanic areas<br />
2. M. alabastrites (p. 110)<br />
21b Apo<strong>the</strong>cia usually dispersed and usually with some hint <strong>of</strong> pigmentation. Spores<br />
(l-)3(-5)-septate, (ll-)15-23(-24)x3-5(-6) /am. Shade forms, occurring on bark <strong>of</strong><br />
old trees (especially Quercus) or on rocks , <strong>of</strong> this widespread and variable species <strong>of</strong> various<br />
substrata (see 27a)<br />
21<br />
22<br />
33. M. peliocarpa (p. 169)<br />
22a (20b) Thallus <strong>of</strong> fragile (easily broken with point <strong>of</strong> dissecting needle) , ash-grey to grey-brown<br />
granular-areolae, <strong>of</strong>ten dissolving to form sorediate patches, PD-I- yellow or red. Usually<br />
sterile<br />
22b Thallus <strong>of</strong> firm areolae or granules, or endoxylic, PD- or PD4- red (M. lignaria). Apo<strong>the</strong>cia<br />
usually numerous<br />
23a (22a) Thallus C-l- red, PD-I- red (gyrophoric acid and argopsin). Spores (l-)3-septate,<br />
14-26(-29)x4-5-5/Ltm<br />
18. M. leprosula (p. 140)<br />
23b Thallus Cf+ red, KC-I- red, PD+ deep yellow (alectorialic acid). Spores 3-7(-9)-septate,<br />
35-45(-60) X 5-6-5 /im<br />
38. M. subleprosula (p. 182)<br />
23<br />
24
104 BRIAN JOHN COPPINS<br />
24a (22b) Thallus PD+ red and C- (argopsin), or PD- and C+ persistent orange (xanthone);<br />
gyrophoric acid absent. Apo<strong>the</strong>cia black, ± globose, sessile or occasionally stipitate. Upper<br />
hymenium green (sometimes brownish in reflexed parts). Hypo<strong>the</strong>cium pale, dilute<br />
greenish or brownish in upper part . Spores 3-7-septate , 16-36(-38) x 4-6(-7) fixn 25<br />
24b Thallus PD— ; thallus and/or apo<strong>the</strong>cia sections C+ red (soon fading; gyrophoric acid) or C—<br />
(no substances) . Apo<strong>the</strong>cia <strong>of</strong>ten adnate at first 26<br />
25a (24a) Areolae C-, PD+ red (argopsin), usually whitish or pale grey. Widespread and<br />
common, especially in upland areas 19a. M. lignaria var. Iignaria(p. 142)<br />
25b Areolae C+ persistent orange, PD— (xanthone), usually whitish with yellowish (isabeUine)<br />
tinge. Local, confined to high rainfall areas, <strong>of</strong>ten with var. lignaria<br />
19b. M. lignaria var. endoleuca (p. 146)<br />
26a (24b) Spores (3-)5-7-septate, (19-)23-34(-34(-38)x4-5-6 fxm. Macroconidia filiform, flexuose,<br />
50-1 10x1 jxm. Thallus and apo<strong>the</strong>cia sections C+ red (gyrophoric acid)<br />
7. M. cinerea (p. 121)<br />
26b Spores mostly 3-septate (5-septate spores absent or in very low frequency), less than 25 (xm<br />
long. Macroconidia less than 50 /i-m long. Thallus and apo<strong>the</strong>cia sections C+ red or C- 27<br />
27a (26b) Thallus and apo<strong>the</strong>cia sections C + red (gyrophoric acid) . Hypo<strong>the</strong>cium usually hyaline<br />
Microconidia (5-)6-7(-7-7)x0-4-0-7 /xm; mesoconidia unknown; macroconidia curved,<br />
27b<br />
21^0(^7) X 1-1 -5 fxm. Widespread on various substrata<br />
Thallus and apo<strong>the</strong>cia sections C— (no substances with t.l.c).<br />
33. M. peliocarpa (p. 169)<br />
Hypo<strong>the</strong>cium with greenbrown<br />
tinge (as in M. lignaria). Microconidia and macroconidia unknown; mesoconidia<br />
4-6-6-3-1 -2-1 -7 fxm. On turf in <strong>the</strong> Arctic; possibly occurring in western Britain on rocks in<br />
coastal districts as a form with a ± obsolete thallus 43. M. ternaria (p. 190)<br />
28a (15b) Thallus <strong>of</strong> well developed, whitish to grey-brown, convex granular-areolae, 0-08-<br />
0-4 mm diam, with intermixed brown cephalodia containing Nostoc or Stigonema. Apo<strong>the</strong>cia<br />
black, convex to subglobose, never markedly constricted below. Spores simple, or<br />
l(-2)-septate spores intermixed, 10-17(-19)x3-5 fxm. Upper hymenium green, K-.<br />
Hypo<strong>the</strong>cium dark. Over bryophytes, plant debris or light soil amongst rocks or on exposed<br />
turf in montane or Arctic regions 29<br />
28b Without <strong>the</strong> above combination <strong>of</strong> characters ; never with cephalodia 30<br />
29a (28a) Hypo<strong>the</strong>cium reddish brown, K— , HNO3-I- bright orange-brown (no purpUsh tinge).<br />
Thallus usually grey-brown. Spores simple or <strong>of</strong>ten 1-septate (rarely 2-septate)<br />
16. M. incrassata (p. 137)<br />
29b Hypo<strong>the</strong>cium purple-brown, K-l- purple intensifying, HNO3-I- purple-red. Thallus usually<br />
whitish or brownish white . Spores mostly simple , septate spores very rare<br />
[if young apo<strong>the</strong>cia turbinate and thallus finely granular, see 51b]<br />
4. M. assimilata (p. 114)<br />
30a (28b) Apo<strong>the</strong>cia whitish, pallid or dull reddish, without distinct pigmentation in section.<br />
Spores mostly 1-septate and over 9 /xm long. Phycobiont micareoid. On bark or lignum or<br />
over bryophytes <strong>the</strong>reon 31<br />
30b Apo<strong>the</strong>cia coloured with obvious pigmentation in section; if not <strong>the</strong>n, spores simple and/or<br />
smaller, or on rock , or phycobiont not micareoid 33<br />
31a (30a) Thallus comprised <strong>of</strong> goniocysts/or apo<strong>the</strong>cia sections C-l- red (gyrophoric acid) and<br />
spores less than 3 fim wide . Shade<br />
forms 8<br />
31b Thallus not comprised <strong>of</strong> goniocysts, endoxylic or weakly areolate, or scurfy granular, or<br />
rimose, sometimes with waxy appearance. All parts C— (gyrophoric acid absent). Spores<br />
<strong>of</strong>ten over 3 fim wide 32<br />
.
LICHEN GENUS MICAREA IN EUROPE 105<br />
32a (31b) Apo<strong>the</strong>cia pale reddish or orange-brown, small, 0-15-0-35 mm diam, sometimes<br />
tuberculate and up to 0-6 mm diam, but never forming large nodulose clusters.<br />
Spores ovoid-fusiform, <strong>of</strong>ten slightly curved, (0-)l(-3)-septate, 9-16x2-5-4-5 /xm. Pale<br />
reddish brown, stalked pycnidia <strong>of</strong>ten present. Thallus usually poorly developed, never<br />
waxy 3. M. anterior (p. 112)<br />
32b Apo<strong>the</strong>cia pallid, at first ± plane and adnate, 0-2-0-4(-0-6) mm diam, later <strong>of</strong>ten coalescing to<br />
form large nodulose clusters up to 2 mm wide. Spores ellipsoid, ovoid or oblong, never<br />
curved, (0-)l -septate, 9-16x3-5 ixm. Without stalked pycnidia, but white, convex sporodochia<br />
(0-1-0-25 mm diam) usually present, bearing simple, ellipsoid macroconidia,<br />
6-10x2-3 /um l.M. adnata (p. 108)<br />
33a (30b) Thallus and apo<strong>the</strong>cia sections C-l- red (? gyrophoric acid). Epi<strong>the</strong>cium fuscous-brown.<br />
Hypo<strong>the</strong>cium pale. Spores markedly curved, 1-septate , 9-12 x 2-5^ fim . On rock<br />
10. M. curvata (p. 126)<br />
33b Thallus and apo<strong>the</strong>cia sections C—; without remaining combination <strong>of</strong> characters 34<br />
34a (33b) Upper hymenium with well defined fuscous-brown epi<strong>the</strong>cium, pigment unchanged in<br />
colour but dissolving into solutions in K. Hypo<strong>the</strong>cium hyaline to dilute yellowish brown.<br />
Phycobiont micareoid. Mostly on lignum or old bark 35<br />
34b Upper hymenium o<strong>the</strong>rwise; if fuscous-brown <strong>the</strong>n pigment not dissolving in K and plant not<br />
on bark or lignum. Hypo<strong>the</strong>cium hyaline, pale or dark. Phycobiont micareoid or not.<br />
Substrata various 36<br />
35a (34a) Spores 0-l(-3)-septate, (9-)ll-15(-19)x 2-3-5 /am. Thallus <strong>of</strong> dispersed to contiguous,<br />
whitish, grey-brown or olivaceous-brown, convex to ± globose areolae. On lignum or old<br />
bark, very rarely on rock 12. M. elachista . (p 131<br />
35b Spores 0(-l)-septate, 6-9-5 x 1 -5-2-3 /xm. Thallus endoxyUc. On lignum<br />
36. M. rhabdogena (p. 181)<br />
36a (34b) Hypo<strong>the</strong>cium hyaHne, yellowish, dilute orange-brown or dilute greenish, not blackish<br />
in thick sections 37<br />
36b Hypo<strong>the</strong>cium dark throughout, <strong>of</strong>ten blackish in thick section 42<br />
37a (36a) Apo<strong>the</strong>cia dark brown or black. Spores 0-1-septate, c. 9-14x4—6 ± m. On exposed<br />
rocks 38<br />
37b Apo<strong>the</strong>cia pale; if dark <strong>the</strong>n spores simple and/or smaller. In sheltered situations on shaded<br />
rocks, exposed tree-roots etc 39<br />
38a (37a) Upper hymenium brown. Phycobiont micareoid. Pycnidia usually present, with ei<strong>the</strong>r<br />
helicoid macroconidia or cylindrical microconidia 39. M. subnigrata (p. 183)<br />
38b Upper hymenium green. Phycobiont not micareoid, thick-walled, 7-21 /xm diam. Pycnidia<br />
unknown 17. M. intrusa (p. 138)<br />
39a (37b) Spores small, in range, 4-8-5x1-2-5 /xm, single or 1-septate. Apo<strong>the</strong>cia small, mostly<br />
less than 0-3 mm diam 40<br />
39b Spores larger, c. 7-10x3^-5 fim, simple. Apo<strong>the</strong>cia usually larger, <strong>of</strong>ten more than 0-3 mm<br />
diam 41<br />
40a (39a) Spores mostly 1-septate, 6-8-5x1-5-2-5 /xm. Apo<strong>the</strong>cia orange-brown or reddish<br />
brown . Hypo<strong>the</strong>cium<br />
dilute orange-brown . Phycobiont (?) micareoid , cells 4-7 /am<br />
28. M. myriocarpa (p. 161)<br />
40b Spores simple, 3-6x1-2 /am. Apo<strong>the</strong>cia blue-grey to blackish. Hypo<strong>the</strong>cium dilute green.<br />
Phycobiont cells 5-12(-18)x 3-8 /am, <strong>of</strong>ten in pairs or short chains Psilolechia clavuiifera (p. 201)<br />
41a (39b) Phycobiont not micareoid, cells 5-12 /am diam. Apo<strong>the</strong>cia pallid to black. Upper<br />
hymenium hyaline to dark green or aeruginose. Hypo<strong>the</strong>cium hyaline, or dilute greenish in<br />
upper parts 5. M. bauschiana (p. 117)<br />
41b Phycobiont micareoid, cells 4-8 /am diam. Apo<strong>the</strong>cia pallid, dull yellowish-orange to reddish<br />
brown. Upper hymenium hyaline. Hypo<strong>the</strong>cium straw-yellow to dilute orange-brown<br />
20. M. lithinella (p. 147)<br />
42a (36b) Hypo<strong>the</strong>cium fuscous or ± reddish brown, without distinct purple or greenish tinges in<br />
water or K, HNO3- or -I- bright orange-brown, never -I- purple-red 43<br />
42b Hypo<strong>the</strong>cium with distinct purple or greenish tinge in water and/or K, HN03-F purple-red 50<br />
)
106 BRIAN JOHN COPPINS<br />
43a (42a) Pycnidia conspicuous, black , sessile or stalked 44<br />
43b Pycnidia inconspicuous and immersed, or absent 45<br />
44a (43a) Spores simple, 5-9x2-5-3-8 /xm. Conidia 2-6-3-6X 1-1 -3 fim. On lignum<br />
24. M. melaeniza (p. 155)<br />
44b Spores 0-l(-3)-septate,8-13(-16)x2.3^)u,m. Conidia 3 • 5-4-8 x 1-1-5 />tm. On shaded rocks,<br />
decaying bryophytes, exposed tree roots or loose stones , rarely on bark or lignum<br />
6. M. botryoides (p. 118)<br />
45a (43b) On rocks, loose stones, consolidated soil, dry bryophyte mats or exposed tree roots in<br />
underhangs, or on lignum 46<br />
45b On mineral soil, or over decaying bryophytes or plant debris on <strong>the</strong> ground; never in<br />
underhangs 48<br />
46a (45a) Spores mostly 1-septate, 6-8-5x1-5-2-5 /xm. Hymenium hyaline or tinged orangebrown,<br />
never greenish. Hypo<strong>the</strong>cium never blackish 28. M. myriocarpa (p. 161)<br />
46b Spores simple. Hymenium <strong>of</strong>ten greenish or ± aeruginose. Hypo<strong>the</strong>cium blackish in thick<br />
section 47<br />
47a (46b) On sheltered rocks, rarely on exposed tree roots. Apo<strong>the</strong>cia convex-globose, <strong>of</strong>ten<br />
tuberculate. Spores 6-8(-9)x2-3(-3-4) /xm. Hypo<strong>the</strong>cium c. 120-360 fxm tall. Phycobiont<br />
not micareoid, cells 5-12 /xmdiam 21. M. lutulata (p. 148)<br />
47b On Hgnum, rarely on rock. Apo<strong>the</strong>cia convex-adnate, never tuberculate. Spores 9-12x<br />
4-5 /im. Hypo<strong>the</strong>cium c. 70-120 fim tall. Phycobiont micareoid, cells 4-7 fxm diam<br />
27. M. muhrii (p. 160)<br />
48a (45b) Spores simple , small , 6-9 - 5 x 3-4 /am . Upper hymenium brown without green or purple<br />
tinge in K. On exposed soil in woodland clearings (? sites <strong>of</strong> old bonfires)<br />
32. M. osloensis (p. 169)<br />
48b Spores larger, mostly over 10 ^tm long. Upper hymenium usually with greenish or purple tinge<br />
inK 49<br />
49a (48b) Thallus blackish. Upper hymenium usually with green tinge in K. Spores simple, or<br />
<strong>of</strong>ten becoming 3-septate (10-)12-21(-25)x(3-5-)4-5 /xm. On exposed soil or truf on<br />
mountain summits or exposed ridges, or at lower altitudes in boreal regions 45. M. turfosa (p. 194)<br />
49b Thallus whitish. Upper hymenium with purple tinge in K. Spores 0-1-septate, (7-)9-5-15x<br />
3-4(-4-5)/am. On fine-sandy or argillaceous soil 23. M. melaenida (p. 154)<br />
50a (42b) Thallus terricolous or muscicolous. Spores simple or 1-septate, mostly in range<br />
9-19x3-4-5 /am. Paraphyses numerous, simple or sparingly branched, 1-1-5 /am wide 51<br />
50b Thallus saxicolous, lignicolous or corticolous, not terricolous or if so (anomalous occurrences)<br />
<strong>the</strong>n spores smaller or paraphyses thinner and much branched 52<br />
51a (50a) On fine sandy or argillaceous soils, never muscicolous or on plant debris; at low<br />
altitudes. Apo<strong>the</strong>cia immarginate, convex to hemisphaerical, adnate when young; in<br />
section without greenish pigmentation in K. Spores 1-septate, (7-)9-5-15 x 3-4 /am<br />
23. M. melaenida (p. 154)<br />
51b On bryophytes or plant debris, sometimes spreading on to sandy soil; in Arctic or Alpine<br />
situations. Apo<strong>the</strong>cia, at least when young, marginate, and markedly constricted below,<br />
turbinate or short-stalked; in section (especially excipulum) with green pigmentation<br />
(<strong>of</strong>ten mixed with purple pigmentation) in K. Spores mostly simple, (9-)10-19(-21)x<br />
(2-5-)3-4-5/am 9. M. crassipes (p. 125)<br />
52a (50b) Phycobiont not micareoid, cells 5-12 /am or more in diam. On shaded rocks in humid<br />
places (narrow ravines or woodlands), or in sheltered underhangs on rock, loose stones,<br />
exposed roots or consolidated soil ; rarely on lignum (old fence posts) 53<br />
52b Phycobiont micareoid , cells 4-8 /am diam . Usually on lignum , rarely on shaded rock 56<br />
53a (52a) Hypo<strong>the</strong>cium dark reddish brown, K- or K± red intensifying, but not turning<br />
purplish or greenish. Hymenium hyaline, dilute green to aeruginose. Spores 6-8(-9)x<br />
2-3(-3-4)/am 21. M. lutulata (p. 148)<br />
53b Hypo<strong>the</strong>cium distinctly greenish in K, more rarely purplish in part or in whole 54
LICHEN GENUS MICAREA IN EUROPE 107<br />
54a (53b) Spores ellipsoid or ovoid-ellipsoid, (6-)7-10x(2-3-)3-4-5 fxm. Apo<strong>the</strong>cia 0-2-0-5 mm<br />
diam, or reaching 1-2 mm when tuberculate. Conidia 3-8-6(-6-6) x 1-1 •7(-2) fjivn<br />
41.M. syIvicoIa(p. 186)<br />
54b Spores ovoid, oblong-ovoid or dacryoid, never ellipsoid, smaller, mostly less than 8 /u,m long<br />
and 2-4 fxm wide. Non-tuberculate apo<strong>the</strong>cia usually smaller, rarely exceeding 0-3 mm<br />
diam. Conidia, if present, shorter 55<br />
55a (54b) Spores 0-1-septate, 5-8(-9) x 1 •5-2-5 fxm. Hypo<strong>the</strong>cium much darker than hymenium.<br />
Pycnidia <strong>of</strong>ten present, 60-120 ^tm diam; wall dark green. Conidia 3-4-3x 1-1-4 fxm.<br />
Phycobiont cells ± globose, c. 5-12 /xm diam, never in short chains.. 44. M. tuberculata (p. 192)<br />
[if similar but phycobiont micareoid, see 58a]<br />
55b Spores simple, 3-6xl-l-7(-2) fim. Hypo<strong>the</strong>cium concolorous with, or paler than, hymenium.<br />
Pycnidia unknown. Phycobiont cells ± globose, eUipsoid or oblong, 5-12(-18)x<br />
3-8 fim, <strong>of</strong>ten in pairs or short chains Psilolechia clavulifera (p. 201)<br />
56a (52b) Spores simple, 6-5-10x2-5-3-7 /xm. Sessile or stalked, black pycnidia present.<br />
Hypo<strong>the</strong>cium purple-brown, K-l- green 29. M. nigella (p. 163)<br />
56b Spores (0-)l -septate. Pycnidia innate, never stalked 57<br />
57a (56b) Spores narrow, oblong-fusiform, 9-14(-16)x 1-8-2-5 /u.m. Hypo<strong>the</strong>cium purple-brown,<br />
K-l- green. Hymenium bright green in upper part. HyaHne paraphyses ra<strong>the</strong>r scanty, thin,<br />
0-7-0-8 /Am wide. Pycnidia with mesoconidia3-9-5-5xl-l-4/Ltm 13. M. exima (p. 134)<br />
57b Spores broader , 2 - 5 fim or more in width , oblong with ± rounded apices , or ovoid 58<br />
58a (57b) Hyahne paraphyses numerous, 1-1-2 ^tm wide. Hypo<strong>the</strong>cium dark sordid-olivaceous or<br />
ohve-brown, without any purple tinge in water or K. Hymenium oHvaceous green. Spores<br />
oblong or ovoid-oblong, (0-)l -septate, (7-)9-12-3x 2-5-3-5 /u,m. Pycnidia with mesoconidia3-4-2x<br />
1-1-3 /xm 31. M. olivacea (p. 167)<br />
58b Hyaline paraphyses scanty or numerous, thin, 0-6-1 /xm wide. Hypo<strong>the</strong>cium <strong>of</strong>ten with purple<br />
tinge in water or K, green and purple pigments <strong>of</strong>ten intermixed 59<br />
59a (58b) Spores ovoid or oblong ovoid, 1-septate, 7-13(-14)x(2-3-)3^-5 fim. Apo<strong>the</strong>cia<br />
0-1-0-2 /xm diam, or reaching 0-3 mm when tuberculate. Thallus endoxylic. Microconidia<br />
4-5x0-8-1 /am; mesoconidia3-8^-7x 1-3-1-8 />tm;macroconidia unknown<br />
8. M. contexta (p. 124)<br />
59b Spores oblong or ovoid oblong, at first 1-septate, becoming 3-septate at maturity. Apo<strong>the</strong>cia<br />
0-12-0-4 mm diam, or reaching 0-5 mm when tuberculate. Thallus superficial, containing<br />
gyrophoric acid when well developed. Microconidia 4-8-7x0-8-1 /xm; mesoconidia unknown;<br />
macroconidia curved, 18-33 x 1-1 -5 /im. Immature forms 22. M. melaena (p. 150)<br />
Key to European species that may occur without apo<strong>the</strong>cia<br />
Notes: all species included here have micareoid algae; all species with stalked pycnidia are<br />
included here.<br />
la Thallus <strong>of</strong> fragile ash-grey or grey-brown granular-areolae <strong>of</strong>ten dissolving or eroding to form<br />
sorediate patches; PD -I- yellow or red . Pycnidia absent 2<br />
lb Thallus o<strong>the</strong>rwise , PD - . Pycnidia usually present and conspicuous ( x 20 lens) 3<br />
2a (la) Thallus C-l- red, PD -I- red (gyrophoric acid and argopsin) 18. M. leprosula (p. 140)<br />
2b Thallus C+ red (<strong>of</strong>ten faint), KC-I- red, PD+ deep yellow (alectorialic acid)<br />
38. M. subleprosula (p. 182)<br />
3a (lb) With stalked or sessile pycnidia containing mesoconidia (smaller ± immersed pycnidia<br />
containing microconidia sometimes present) ; never with curved or flexuose macroconidia 4<br />
3b Pycnidia innate sometimes becoming emergent with gaping ostioles, or with small sessile<br />
apo<strong>the</strong>cia-Hke sporodochia; pycnidia sometimes containing curved or flexuose macroconi-<br />
dia<br />
4a (3a) Pycnidia black<br />
-^<br />
4b Pycnidia whitish to reddish brown, never black °<br />
5a (4a) Pycnidial wall oHvaceous brown, K-l- violet. Mesoconidia 3-5-5 x 1-1 -5(-l -7) fim<br />
5b Pycnidial wall K- or K-l- olivaceous<br />
11<br />
26. M. misella (p. 158)<br />
"
108 BRIAN JOHN COPPINS<br />
6a (5b) Pycnidial wall and stalk dark purple-brown, K-l- dark green. Mesoconidia 3-4-4-3X<br />
l-2-l-6)u,m 29. M. nigella (p. 163)<br />
6b Pycnidial wall dark olivaceous, K— ; stalk fuscous or reddish-brown, K— 7<br />
7a (6b) Mesoconidia 3-5-4-8X 1-1-5 /xm. On bryophytes, shaded rocks and stones or exposed<br />
roots, rarely on lignum 6. M. botryoides (p. 118)<br />
7b Mesoconidia2-6-3-6xl-l-3/Lim. Onlignum 24. M. melaeniza (p. 155)<br />
8a (4b) Thallus <strong>of</strong> pale to dark green goniocysts, 12-40 /x.m diam; goniocysts containing purple<br />
oily substance in K. Pycnidia brown with white tomentum. Mesoconidia (4-)4-5(-6)x<br />
1-3-1-7/xm 15. M. hedlundii (p. 135)<br />
8b Thallus indistinctly areolate or rimose, or endoxylic, without goniocysts, K— . Pycnidia<br />
without tomentum 9<br />
9a (8b) Pycnidia whitish, C-l- red (gyrophoric acid) . Mesoconidia<br />
4—6 x 1-1-5 /im<br />
35. M. pycnidiophora (p. 179)<br />
9b Pycnidia C— , whitish or reddish brown 10<br />
10a (9b) Pycnidia whitish, <strong>of</strong>ten branched. Mesoconidia 6-8x1-1-8 ^tm. Usually on bark, in<br />
oceanic areas 37. M. stipitata (p. 182)<br />
10b Pycnidia reddish brown, rarely branched. Mesoconidia 3-5-4-5 x 1-2-1-6 fim. On lignum in<br />
boreal regions 3. M. anterior (p. 112)<br />
11a (3b) With whitish, cushion-like, apo<strong>the</strong>cia-like sporodochia c. 0-1-0-25 mm diam, bearing<br />
oblong-ellipsoid macroconidia 6-10x2-3 /xm (immersed, inconspicuous pycnidia containing<br />
mesoconidia 4-5-3x1-2-1-5 /am may also be present). All parts C- (t.l.c: nil). On<br />
lignum or decaying bark <strong>of</strong> old stumps 1. M. adnata (p. 108)<br />
lib Sporodochia absent . Thallus and/or pycnidia C -I- red (gyrophoric acid) or C - (t . 1 . c . : 'prasina<br />
unknowns') 12<br />
12a (lib) Thallus <strong>of</strong> pale to dark green (sometimes K-l- violet) goniocysts 12-60 /x,m; pycnidia, if<br />
present, 30-80 yam diam containing ei<strong>the</strong>r (5-)5 -5-8x0-7-1 /xm microconidia, or<br />
(3-5-)4-6 X 1-2-1-7 ^tm mesoconidia. All parts C- (t.l.c. : 'prasina unknowns')<br />
34. M. prasina (p. 173)<br />
12b Thallus areolate, or scurfy (invaded by dematiaceous hyphae and foreign algae), or ±<br />
endoxylic; thallus and/or pycnidia usually C-l- red (gyrophoric acid) ; sometimes with curved<br />
or flexuose macroconidia 13<br />
13a (12b) With numerous pycnidia containing mesoconidia, 3-5-4-5x1-3-2 /xm, <strong>of</strong>ten<br />
extruded as conspicuous white blobs; in addition, pycnidia containing microconidia<br />
(4-)4-5-6(-6-5)x 0-7-1 /xm, or curved macroconidia 12-12x1 /xm, sometimes present;<br />
pycnidia walls with pale olivaceous pigment, K-l- violet. Usually on lignum, commonly on<br />
worked timber 11. M. denigrata(p. 127)<br />
13b Pycnidia usually with longer, curved or flexuose macroconidia; smaller pycnidia with microconidia<br />
sometimes present; mesoconidia unknown; pycnidial walls hyaline or greenish, K—<br />
On various substrata, rarely on worked timber 14<br />
14a (13b) Macroconidia curved or hamate, 21-40x1-1-5 /xm. Microconidia (5-)6-7(-7-7)x<br />
0-4-0-7 IX,. On various substrata but most commonly encountered without apo<strong>the</strong>cia on<br />
bark <strong>of</strong> old trees (especially Quercus) in old woodlands 33. M. peliocarpa (p. 169)<br />
14b Macroconidia flexuose, 50-110x1 ixm. Microconidia (3-8-)4-5x0-5-0-7 /xm. Seen without<br />
apo<strong>the</strong>cia on shaded lignum in woodland, and over bryophytes in open montane situations<br />
1. Micarea adnata Coppins, sp. nov.<br />
(Figs 7A, 36-37; Map 1)<br />
The species<br />
7. M. cinerea (p. 121)<br />
Thallus effusus, tenuis, ± laevis vel furfuraceo-granulosus, pallide griseo-viridis, plerumque nonnihil<br />
cereus. Algae cellulis 4—7 fxm diam. Apo<strong>the</strong>cia pallide eburnea vel straminea vel demum pallide rufescen-<br />
tia, immarginata, adnata, primum planiuscula mox convexa, 0-2-0-4(-0-6) mm diam, interdum coalescentia<br />
in fasciculosos noduliformes ad 2 mm diam. Hymenium 35-40 ^im altum, ± hyalinum. Ascosporae<br />
ellipsoideae, ovoideae, oblongo-ellipsoideae, oblongo-ovoideae vel oblongae, (O-)l-septatae, 9-16x<br />
.
LICHEN GENUS MICAREA IN EUROPE 109<br />
3-5 />tm. Paraphyses numerosae, ramosae et anastomosantes, c. 1-1-5 jxm latae, apicibus vix incrassatis.<br />
Hypo<strong>the</strong>cium hyalinum. Excipulum bene evolutum et manifestum praesertim in apo<strong>the</strong>ciis junioribus.<br />
Conidiomata pycnidiiformia et sporodochiiformia. Pycnidia pauca et inconspicua, ± immersa, alba, c.<br />
40-60 /xm diam, producentia conidiis cylindricis 4-5-3x 1-2-1-5 ^tm. Sporodochia plerumque numerosa et<br />
conspicua, alba, pulvinata, 20-250 jxm diam, producentia conidiis cylindricis vel oblongo-ellipsoideis<br />
6-5-9-5X 2-3-3 ^tm. Thallus et apo<strong>the</strong>cia, K— , C—<br />
, PD-; sine materia chemica.<br />
Typus: Caledonia, Argyll, Dunoon, Benmore, ad River Eachaig, ad corticem Alni, 18 xi 1977, leg. B. J.<br />
Coppins 3256 (E-holotypus).<br />
Thallus effuse, thin, ± smooth or finely scurfy-granular, pale grey-green, ra<strong>the</strong>r waxy in<br />
appearance; white, arachnoid, prothalline hyphae sometimes visible (x50). Phycobiont<br />
micareoid, cells 4-7 jxm diam.<br />
Apo<strong>the</strong>cia usually numerous but occasionally few or absent, pallid to pale straw-brown or pale<br />
reddish brown; at first ± plane, adnate, ± immarginate but <strong>of</strong>ten with a white rim, c. 30-50 /am<br />
wide, from which a few white, arachnoid hyphae are sometimes seen to arise and penetrate into<br />
<strong>the</strong> thallus; later becoming convex although remaining adnate and never becoming constricted<br />
below; dispersed, or <strong>of</strong>ten confluent, and frequently coalescing to form large, irregular, waxy,<br />
nodulose clusters; individual apo<strong>the</strong>cia 0-2-0-4(-0-6) mm diam, clusters up to 2 mm broad.<br />
Hymenium 35-40 /xm tall, hyaHne or dilute straw-yellow, K-. Asci clavate, 30-35x10-12 /xm.<br />
Spores ellipsoid, ovoid, oblong-ellipsoid, oblong-ovoid or oblong, <strong>of</strong>ten slightly constricted at<br />
<strong>the</strong> septum, (0-)l -septate, 9-16x3-5 fxm. Paraphyses numerous, branched and anastomosing,<br />
c. 1-1-5 fjLm wide and scarcely widening above; apices much branched and entangled. Hypo<strong>the</strong>cium<br />
c. 40-120 /xm tall, hyaline; hyphae hyaline, c. 1-2 ^tm wide, interwoven but becoming ±<br />
vertically orientated towards <strong>the</strong> hymenium, intermixed with short-celled ascogenous hyphae<br />
that are up to 5 /xm wide. Excipulum well developed and clearly seen in young apo<strong>the</strong>cia, c.<br />
30-35 /xm wide; hyphae radiating, branched and anastomozing, c. 1-2-5 /xm wide.<br />
Conidiomata <strong>of</strong> two types, one pycnidial, <strong>the</strong> o<strong>the</strong>r sporodochial. Pycnidia sometimes present<br />
but <strong>the</strong>n usually few in number and inconspicuous, white, immersed, c. 40-60 ^im diam;<br />
conidiogenous cells ± cylindrical, 6-10x1-1-5 /xm; conidia (mesoconidia) cylindrical, sometimes<br />
faintly biguttulate, 4—5-3xl-2-l-5 /xm. Sporodochia usually present, numerous and<br />
conspicuous, resembling small apo<strong>the</strong>cia, white or pallid, pulvinate, 80-250 /xm diam; conidiogenous<br />
cells cylindrical, 10-20x1-7-2 fxm; conidia (macroconidia) cylindrical to oblong<br />
ellipsoid or occasionally oblong-obovoid, simple, eguttulate, 6-5-9-5x2-3-3 fxm.<br />
Chemistry: Thallus and apo<strong>the</strong>cia K— , C— , PD-; no substances detected by t.l.c.<br />
Observations: Micarea adnata is easily recognised by <strong>the</strong> combination <strong>of</strong> a pale grey-green, ±<br />
waxy thallus, pale, adnate to clustered apo<strong>the</strong>cia and, above all, by <strong>the</strong> presence <strong>of</strong> sporodochia.<br />
These cushion-like conidiomata, which resemble small apo<strong>the</strong>cia, produce conidia externally<br />
over <strong>the</strong>ir surface; such structures are unknown in any o<strong>the</strong>r Micarea, and appear not to have<br />
been previously reported for lichenised fungi. The ra<strong>the</strong>r large, non-septate, oblong conidia<br />
produced by <strong>the</strong> sporodochia are probably homologous to <strong>the</strong> curved or filiform and <strong>of</strong>ten<br />
septate macroconidia <strong>of</strong> such species as M. cinerea, M. denigrata, and M. peliocarpa. Some<br />
specimens <strong>of</strong> M. adnata have a few pycnidia with typical mesoconidia, but microconidia have not<br />
yet been found.<br />
It is curious that this most distinctive species had not previously been afforded taxonomic<br />
recognition, although it was well known by Arnold who erroneously referred it to Biatorina<br />
prasiniza, a synonym <strong>of</strong> M. prasina. It is with forms <strong>of</strong> M. prasina with pallid apo<strong>the</strong>cia that M.<br />
adnata is most likely to be confused. However, <strong>the</strong> former has a non-waxy, granular thallus<br />
composed <strong>of</strong> goniocysts, apo<strong>the</strong>cia which soon become constricted below, usually smaller<br />
spores and an apparent absence <strong>of</strong> macroconidia. M. prasina is sometimes found to have<br />
pycnidia containing mesoconidia similar to those <strong>of</strong> M. adnata, but more frequently it has<br />
pycnidia containing microconidia. In addition, well developed specimens oiM. prasina have one<br />
<strong>of</strong> three distinctive substances detectable by t.l.c, whereas M. adnata contains no lichen<br />
substances. M. anterior is similar to M. adnata in being lignicolous with pale apo<strong>the</strong>cia and<br />
spores that are mainly 1-septate; however, its apo<strong>the</strong>cia are usually darker, varying from a pale
110 BRIAN JOHN COPPINS<br />
reddish or orange-brown to red-brown, with spores that are relatively narrower and ± fusiform,<br />
and a reddish brown excipulum. Fur<strong>the</strong>rmore, M. anterior has its mesoconidia borne in reddish<br />
brown, stalked pycnidia.<br />
Habitat and distribution: M. adnata is usually found on old stumps and associated decaying<br />
bryophyte mats and peaty debris; <strong>the</strong> Ugnum <strong>of</strong> such stumps is usually s<strong>of</strong>t and riddled with holes<br />
bored by insects. In addition, it is found on <strong>the</strong> loose bark <strong>of</strong> trunks <strong>of</strong> old trees, including ^/nwj^,<br />
Betula, Corylus, Quercus, Abies, and Pinus. It usually occurs in extensive patches with few<br />
associated lichens, but amongst those noted are Cladonia spp. (including C. coniocraea and C.<br />
squamosa), Micarea prasina, Parmelia laevigata, and Platismatia glauca. Most collections are<br />
from old, ± natural deciduous or coniferous woodlands, but in western Scotland it is found also<br />
in old estate woodlands (possibly created from former ± natural woodland) containing many<br />
exotic trees.<br />
The distribution <strong>of</strong> M. adnata in Britain is centred on western Scotland, with a few outlying<br />
localities in Cumbria and north Wales. It is so far unrecorded for south-west England or western<br />
Ireland, and it should be sought for in <strong>the</strong> old oak-woods <strong>of</strong> those regions. Its distribution<br />
coincides with an annual rainfall <strong>of</strong> over 1000 mm per year, distributed over at least 160 rain<br />
days, and is attributable to <strong>the</strong> General Western group <strong>of</strong> Coppins (1976). Similarly, many <strong>of</strong> its<br />
localities on mainland Europe are in regions with a high annual precipitation (over 1000 mm per<br />
year), viz. sou<strong>the</strong>rn Bavaria, Austria (Steiermark), and Switzerland (Bern). However, <strong>the</strong>re are<br />
a few specimens from lower rainfall districts, namely, middle and nor<strong>the</strong>rn Bavaria, and eastern<br />
Sweden (Sodermanland), perhaps in situations where local topographical or vegetational<br />
features provide a microclimate sufficient to compensate for <strong>the</strong> low rainfall. If <strong>the</strong> distribution<br />
<strong>of</strong> M. adnata is largely controlled by microclimatic conditions, usually resulting from a high<br />
rainfall, <strong>the</strong>n it should be expected to extend to Norway, at least <strong>the</strong> south-western part <strong>of</strong> that<br />
country.<br />
Exsiccata: Arnold Lich. Mon. 244 (BM ex K, M). Britzelmayer Lich. Exs. 174 (M).<br />
2. Micarea alabastrites (Nyl.) Coppins<br />
(Figs 2, 6, 8; Map 2)<br />
in Topham & Walker in Lichenologist 14: 66 (1982). -<br />
Lecidea alabastrites Nyl. in Flora, Jena 62: 207 (1879). - Bilimbia sphaeroides var. alabastrites (Nyl.) A.L.<br />
Sm., Monogr. Br. Lich. 2: 138 (1911). Type: Ireland, West Galway, Derryclare, near Kylemore, on<br />
bryophytes on bark, 1878, C. Larbalestier (H-NYL 18656-hoIotype!; BM-isotype!).<br />
Thallus effuse, sometimes partly immersed in <strong>the</strong> substratum (especially when on lignum),<br />
but usually well developed on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> substratum as crowded, confluent, ±<br />
convex-hemispherical to subglobose areolae. Areolae dull whitish, greenish white, or pale dull<br />
green, matt or slightly glossy, c. 40-140 /x,m diam, or up to 250 ^im if containing a pycnidium <strong>of</strong><br />
<strong>the</strong> macroconidial anamorph. Areolae in section, ecorticate but with a hyaline amorphous<br />
covering layer c. 4-10 jxm thick.<br />
Apo<strong>the</strong>cia usually numerous and crowded, sometimes confluent and coalescing in groups <strong>of</strong><br />
up to eight apo<strong>the</strong>cia, adnate, plane to shallow-convex or convex-hemispherical, occasionally<br />
tuberculate, large, shallow-convex apo<strong>the</strong>cia sometimes flexuose, <strong>of</strong>ten with an indistinct,<br />
slightly paler margin that is flush with <strong>the</strong> level <strong>of</strong> <strong>the</strong> disc, whitish, cream-white, ivory-white or<br />
pallid, 0-2-0-7 mm diam, or up to 0-9 mm diam when tuberculate. Hymenium 45-55 yam tall,<br />
hyaline, or dilute straw in upper part. Asci clavate, 42-50x12-16 /xm. Spores fusiform or<br />
clavate-fusiform, sometimes slightly curved, 3(-5)-septate, occasional 7-septate spores encountered,<br />
(16-)18-26(-29)x(4-)4-5-5(-6) jxm. Paraphyses numerous, branched, <strong>of</strong>ten anastomosing,<br />
c. 1-1-5 /xm wide; apices <strong>of</strong>ten more richly branched and entangled, <strong>of</strong>ten slightly<br />
incrassate to T8 /xm. Hypo<strong>the</strong>cium c. 45-100 /am tafl; hyphae interwoven, c. 1-T7 /xm;<br />
ascogenous hyphae with swollen cells c. 2-4 /xm wide. Excipulum well developed (laterally c. 45<br />
/x wide in young apo<strong>the</strong>cia), hyaline, <strong>of</strong> richly branched and anastomosing, radiating hyphae, c.<br />
1-T5 /xm wide.
LICHEN GENUS MICAREA IN EUROPE 111<br />
Map 1 Micarea adnata # 1950 onwards O Before 1950<br />
Pycnidia usually present, white with hyaline walls, <strong>of</strong> two types: (a) immersed in areolae, c.<br />
100-220 /xm diam, ostiole <strong>of</strong>ten widely gaping; conidia (macroconidia) markedly curved, <strong>of</strong>ten<br />
sigmoid, sometimes faintly 3-5-septate, 21-55 xc. 1 (xm; (b) ± sessile, c. 50-100 ^tm diam,<br />
ostiole not, or only slightly gaping; conidia (microconidia) narrowly fusiform-cylindrical,<br />
5-7x0-5-0-7/am.<br />
Chemistry: Thallus and apo<strong>the</strong>cia C-l- red; t.l.c: gyrophoric acid.<br />
Observations: M. alabastrites is morphologically and chemically very similar to M. peliocarpa,<br />
and it could easily be dismissed as representing a pale, shade form <strong>of</strong> that species. Apart from <strong>the</strong><br />
complete absence <strong>of</strong> pigment from all its tissues, M. alabastrites is subtly different from M.<br />
peliocarpa in having slightly larger apo<strong>the</strong>cia and spores. Shade forms <strong>of</strong> M. peliocarpa are<br />
usually sparingly fertile with scattered apo<strong>the</strong>cia, whereas <strong>the</strong> apo<strong>the</strong>cia <strong>of</strong> M. alabastrites are<br />
almost invariably numerous, crowded and <strong>of</strong>ten confluent. My early doubts regarding <strong>the</strong><br />
distinction <strong>of</strong> <strong>the</strong>se two species were dispelled by <strong>the</strong> collection, on several occasions, <strong>of</strong> M.<br />
alabastrites with adjacent thalli <strong>of</strong> M. peliocarpa with blackish apo<strong>the</strong>cia; <strong>the</strong> slight differences in<br />
<strong>the</strong> sizes <strong>of</strong> apo<strong>the</strong>cia and spores were confirmed with <strong>the</strong>se collections. The two species also<br />
differ somewhat in distribution and habitat; M. peliocarpa is widely distributed throughout<br />
much <strong>of</strong> Europe on a wide range <strong>of</strong> substrata (tree trunks, mossy rocks peaty soil etc.); whereas<br />
M. alabastrites has a hyperoceanic distribution and occurs on tree trunks (or on bryophytes<br />
<strong>the</strong>reon) and occasionally lignum, but apparently never on mossy rocks or peaty soil.<br />
In <strong>the</strong> recent checklist <strong>of</strong> <strong>British</strong> lichens (Hawksworth et al, 1980) I mistakenly placed
112 BRIAN JOHN COPPINS<br />
Table 4 Diagnostic features for <strong>the</strong> separation <strong>of</strong> Micarea alabastrites, M. cinerea, and M. peliocarpa.<br />
Apo<strong>the</strong>cia size (mm)<br />
Apo<strong>the</strong>cia colour<br />
Hymenium height (/xm)<br />
Spore septation<br />
Spore length (/xm)<br />
Spore breadth<br />
Macroconidia shape<br />
Macroconidia length (/xm)<br />
Thallus colour<br />
peliocarpa alabastrites cinerea<br />
0.14^0-4(-0-6)<br />
pallid to black<br />
40-55<br />
(l-)3(-5)<br />
(ll-)15-23(-24)<br />
3-5(-6)<br />
curved-sigmoid<br />
21^0(-50)<br />
greenish white to<br />
dark blue-grey<br />
0-2-0-7<br />
whitish or pallid<br />
45-55<br />
3(-7)<br />
(16-)18-26(-29)<br />
(4-)4-5-5(-6)<br />
curved-sigmoid<br />
21-55<br />
greenish white or<br />
pale green<br />
0-2-0-7<br />
pallid to black<br />
55-70<br />
(3-)7<br />
(19-)23-34(-38)<br />
4-5-6<br />
flexuose<br />
50-110<br />
greenish white to<br />
dark blue-grey<br />
'alabastrites' as a synonym <strong>of</strong> M. cinerea. At that time I had only seen <strong>the</strong> fragmentary isotype <strong>of</strong><br />
Lecidea alabastrites in BM; I found this to have a few 7-septate spores and considered it a<br />
juvenile, shade-form <strong>of</strong> M. cinerea. Subsequent examination <strong>of</strong> <strong>the</strong> holotype (a large, healthy<br />
specimen) proved me to be wrong, and that it actually belonged to a hyperoceanic species which<br />
I had intended to describe as new to science. M. cinerea is ano<strong>the</strong>r close relative <strong>of</strong> M.<br />
alabastrites and is more prone to occur with ± pallid apo<strong>the</strong>cia than M. peliocarpa. It differs<br />
from both <strong>the</strong>se species in having spores which are mostly 7-septate at maturity, and macroconi-<br />
dia which are flexuose and much longer; for a comparison <strong>of</strong> <strong>the</strong>se three species see Table 4.<br />
Occasional 7-septate spores have been found in several specimens <strong>of</strong> M. alabastrites, but <strong>the</strong>y<br />
never number more than two or three in each squash preparation.<br />
M. alabastrites is one <strong>of</strong> <strong>the</strong> several lichens, named 'Bacidia (or Bilimbia) sphaeroides' by<br />
<strong>British</strong> lichenologists. This name is based on Lichen sphaeroides Dickson, which is a species <strong>of</strong><br />
Biatora Fr. (non Ach.) and currently known as Catillaria sphaeroides (Dickson) Schuler. The<br />
name Bacidia sphaeroides as commonly used by Scandinavian workers refers to a species (not<br />
known in Britain) which also belongs in Biatora; its correct name (basionym) is probably<br />
Bilimbia tetramera de Not.<br />
Habitat and distribution: In <strong>the</strong> <strong>British</strong> Isles M. alabastrites is mainly found in communities <strong>of</strong><br />
<strong>the</strong> Parmelietum laevigatae association on <strong>the</strong> trunks <strong>of</strong> Quercus, Betula, and, more rarely,<br />
Alnus, Fagus, Crataegus, Ilex, Pinus, Pseudotsuga, and Juniperus. Associated lichens include<br />
Bryoria fuscescens, Catillaria pulverea, Cladonia spp., Hypogymnia physodes, Lepraria incana<br />
agg., Micarea cinerea, M. peliocarpa, M. stipitata, M. syno<strong>the</strong>oides, Mycoblastus sanguinarius,<br />
M. sterilis, Parmelia crinita, P. laevigata, P. saxatilis, Platismatia glauca, Sphaerophorus<br />
globosus, and Usnea spp. Occasionally it is found on lignum <strong>of</strong> fallen decorticate trunks,<br />
especially in <strong>the</strong> western native pinewoods; associated species include Lecidea granulosa agg.,<br />
Micarea lignaria, M. peliocarpa, and Platismatia glauca. To date, it is not known to occur on<br />
mossy boulders, directly on rock, or on <strong>the</strong> ground on peaty debris, etc.<br />
Its distribution in <strong>the</strong> <strong>British</strong> Isles is correlated with areas experiencing at least 180 'wet days'<br />
per annum and is referable to <strong>the</strong> 'General Western Group' <strong>of</strong> Coppins (1976). Elsewhere it is<br />
known from western Norway (Hordaland), <strong>the</strong> Azores (on Cryptomeria), and <strong>the</strong> Canary<br />
Islands {on Erica arborea).<br />
3. Micarea anterior (Nyl.) Hedl.<br />
(Figs7B,38C-D)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 76, 86 (1892). - Lecidea anteriorNyl. in Flora, Jena<br />
58: 299 (1875). - Catillaria anterior (Nyl.) Zahlbr., Cat. lich. univ. 4: 29 (1926). Type: Finland, Tavastia<br />
australis, Asikkala, 1863, /. P. Norrlin (H-NYL 21655 - lectotype!; H - isolectotypes!).<br />
Micarea anterior i. diluta Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 76, 86 (1892). Type:<br />
Sweden, Halsingland, Jarvso, vii 1890,7. T. Hedlund{S-ho\o\ypt\; UPS -isotype!).
LICHEN GENUS MICAREA IN EUROPE 113<br />
Map 2 Micarea alabastrites # 1950 onwards O Before 1950<br />
Thallus effuse, endoxylic, or epixylic, <strong>the</strong>n whitish and irregularly verrucose to 100 /xm thick,<br />
sometimes becoming secondarily rimose, but without distinct primary areolae; in section,<br />
without an amorphous covering layer. Phycobiont micareoid, cells 4-7 ^tm diam.<br />
Apo<strong>the</strong>cia numerous, few, or absent, immarginate, convex, adnate, <strong>of</strong>ten becoming tuberculate,<br />
pale reddish brown, dull orange-brown or red-brown, 0- 15-0-35 mm diam, or to 0-6 mm<br />
when tuberculate, Hymenium 38-50 /zm tall, ± hyahne with irregular, dilute red-brown (K— , or<br />
dulling) blotches, especially in upper part. Asci clavate or cylindrical-clavate, 35^5 x 10-12 /xm.<br />
Spores oblong-ovoid to fusiform or ± bifusiform, <strong>of</strong>ten slightly curved, (0-)l(-2 or ?-3)-septate,<br />
9-14(-16)x2-5-4(-4-5) jxm. Paraphyses ra<strong>the</strong>r numerous, simple or sparingly branched, many<br />
becoming much branched above with <strong>the</strong>ir apices entangled, 0-6-1 /xm wide; apices <strong>of</strong>ten<br />
irregularly incrassate, hyaline and up to l-5(-2) ^im wide, or sometimes thickened with reddish<br />
brown pigment and up to 3 ^tm wide. Hypo<strong>the</strong>cium c. 60-110 ^lm. tall, hyaline. Excipulum thin<br />
but clearly seen in sections as a reddish brown reflexed zone; hyphae radiating, branched and<br />
anastomosing, hyaline and c. 1 fxm wide, but becoming thickened with brownish pigment and up<br />
to 2 jxm wide in <strong>the</strong> outer excipulum.<br />
Pycnidia usually numerous; <strong>of</strong> two types: (a) sessile on old apo<strong>the</strong>cia, but more usually borne<br />
on slender stalks (pycnidiophores) which arise from <strong>the</strong> thallus or in some cases from old<br />
apo<strong>the</strong>cia; 70-250 yam tall and 40-70 ^tm diam (overall); stalks pallid but <strong>of</strong>ten with reddish<br />
brown blotches, glabrous, simple, or sometimes branched and bearing two or three pycnidia;<br />
pycnidia (excluding stalk) short cylindrical or slightly tapering inwards above, 40-80 /xm tall and<br />
40-70 /xm diam, with dark reddish brown walls, K- or duUing; conidia {mesoconidia) cylindric-
114 BRIAN JOHN COPPINS<br />
al, <strong>of</strong>ten faintly biguttulate and slightly constricted in <strong>the</strong> middle, 3-5-4-5xl-2-l-6 /xm; (b)<br />
sessile on <strong>the</strong> thallus or old apo<strong>the</strong>cia, c. 30-40 /xm diam, with dark reddish brown walls, K— or<br />
dulling; conidia (microconidia) narrowly cylindrical, (3-2-)3-6-4-5x0-7-0-9 /xm.<br />
Chemistry: Thallus K— , C—<br />
detected by t. I.e.<br />
, PD—<br />
; sections <strong>of</strong> thallus and apo<strong>the</strong>cia C-; no substances<br />
Observations: Micarea anterior is characterised by its reddish brown apo<strong>the</strong>cia, ra<strong>the</strong>r<br />
narrow, mostly 1-septate spores, pale pycnidiophores bearing dark brown pycnidia, and a ±<br />
endoxylic, or thin and irregularly verrucose, thallus. Stalked pycnidia are a feature <strong>of</strong> several<br />
o<strong>the</strong>r hgnicolous species (e.g. M. melaeniza, M. misella, M. nigella, and <strong>the</strong> occasionally<br />
lignicolous M. botryaides), but in those species <strong>the</strong> pycnidiophores appear black and have<br />
different pigmentations when examined microscopically. The pycnidiophores <strong>of</strong> M. hedlundii<br />
are brownish but distinctly tomentose, and those <strong>of</strong> <strong>the</strong> normally corticolous M. pycnidiophora<br />
and M. stipitata are entirely hyaline. M. adnata has its mesoconidia produced in immersed<br />
pycnidia and has macroconidia borne on small apo<strong>the</strong>cia-like sporodochia; fur<strong>the</strong>r distinguishing<br />
features from M. anterior include a ± waxy appearance <strong>of</strong> <strong>the</strong> whole plant, mostly<br />
paler apo<strong>the</strong>cia, and relatively broader spores with rounded apices. Richly fertile specimens <strong>of</strong><br />
M. anterior could be confused with Catillaria erysiboides {q. v. ) but that species has marginate<br />
young apo<strong>the</strong>cia, short ovoid spores, and a non-micareoid phycobiont.<br />
Habitat and distribution: M. anterior is found on <strong>the</strong> ra<strong>the</strong>r s<strong>of</strong>t and smooth lignum <strong>of</strong><br />
decorticate trunks <strong>of</strong> Picea and Pinus, and associated with such species as Cetraria pinastri,<br />
Cladonia spp., Hypogymnia physodes, Lecidea icmalea, Lepraria incana agg., Micarea adnata,<br />
M. contexta, M. misella, M. prasina, Parmeliopsis ambigua, P. hyperopta, Pertusaria pupillaris,<br />
and <strong>the</strong> non-lichenised Cryptodiscus pallidas. It is a rare or overlooked species, apparently<br />
confined to Scandinavia, from where it is known from several locahties in middle and sou<strong>the</strong>rn<br />
Sweden, and south-west Finland.<br />
Exsiccata: Malme Lich. Suec. 22 (M, S).<br />
4. M. assimilata (Nyl.) Coppins, comb. nov.<br />
(Fig. 9; Map 3)<br />
Lecidea assimilata Nyl., Lich Scand.: 221 (1861). Type: Norway, Nordland, Helgeland, M. N. Blytt<br />
(H-NYL 16556- lectotype!).<br />
Lecidea assimilata a. [var.] irrubata Th. Fr., Lich. Scand. 2: 522 (1874); nom. inval. (Art. 26).<br />
Thallus growing on bryophytes, plant debris or sandy soil, composed <strong>of</strong> confluent, irregular,<br />
convex-verrucose, sometimes flattened and subeffigurate areolae that are <strong>of</strong>ten intermixed with<br />
cephalodia. Areolae white or brownish white, matt or sometimes glossy, 0-08-0-4 mm diam; in<br />
section, sometimes with a hyaline amorphous covering layer up to 7 /xm thick, outermost hyphae<br />
hyahne; a ± algal-free medulla sometimes differentiated in large areolae. Thallus hyphae c.<br />
T8-3 /xm diam. Phycobiont micareoid, cells 4—7 fxm diam. Cephalodia <strong>of</strong>ten present, irregularly<br />
globose and hidden amongst <strong>the</strong> areolae, sometimes visible externally as brown areolae-like<br />
structures, 0-2-0-4 mm diam; containing Nostoc, cells 3-5 fxm diam. Less <strong>of</strong>ten present are<br />
irregular, ra<strong>the</strong>r loose clusters (? cephalodia) <strong>of</strong> Stigonema.<br />
Apo<strong>the</strong>cia numerous, immarginate, convex to subglobose, matt or ± glossy (subnitid),<br />
0-3-0-8 mm diam, sometimes forming tuberculate clusters up to T5 mm diam. Hymenium,<br />
45-50 /xm tall; upper part (epi<strong>the</strong>cium) dark aeruginose, olivaceous or brownish-green, K—<br />
HNO3+ red; mid-hymenium dilute greenish; lower hymenium dilute greenish and K— , or dilute<br />
purplish and K-l- purple intensifying or K-l- sordid green. Asci clavate, 45-50x 12-14 /xm. Spores<br />
oblong-ellipsoid to oblong-fusiform, 0(-l)-septate, (10-)12-16(-19)x3-5 /xm. Paraphyses<br />
numerous, simple below, but <strong>of</strong>ten forked or branched above, 1-5-1 •7(-2) jxm wide, sometimes<br />
widening above to 3 /xm; apical walls hyaline although surrounded by densely pigmented<br />
gel-matrix. Hypo<strong>the</strong>cium c. 150-400 /xm tall, dark purple brown, K-t- purple intensifying or<br />
(especially in upper part) K-l- dark green; all parts HNO3-I- purple-red; hyphae interwoven but<br />
,
LICHEN GENUS MICAREA IN EUROPE 115<br />
± vertically orientated near <strong>the</strong> hymenium, c. 1-7-2-7 fxm wide, embedded in a densely<br />
pigmented matrix; ascogenous hyphae c. 1-7-2-7 /xm wide, embedded in a densely pigmented<br />
matrix; ascogenous hyphae c. 2-5-5 fxm wide. Excipulum reflexed, mottled reddish brown or<br />
sordid olivaceous; hyphae radiating, branched and anastomosing, 1-5-2 /xm wide.<br />
Pycnidia rare, semi-immersed to sessile, black, 40-100 /^m diam; walls dull reddish brown, or<br />
dull olivaceous in part; conidia (? microconidia) cylindrical, 6-9x1-1-5 /xm.<br />
Chemistry: Thallus K— , C— , KC—<br />
, PD — ; t.l.c: no substances.<br />
Observations: Micarea assimilata is characterized by its conspicuous, whitish, verrucose<br />
areolae which are <strong>of</strong>ten intermixed with brown cephalodia, ra<strong>the</strong>r large, convex, black<br />
apo<strong>the</strong>cia, green (K— ) epi<strong>the</strong>cium, dark purple-brown hypo<strong>the</strong>cium and predominantly simple,<br />
large spores. It is apparently closely related to M. crassipes, M. incrassata, M. melaenida, and M.<br />
subviolascens , all <strong>of</strong> which share ra<strong>the</strong>r stout, simple or sparingly branched paraphyses, ra<strong>the</strong>r<br />
stout excipular hyphae, dark hypo<strong>the</strong>cia and an absence <strong>of</strong> lichen substances. In many ways M.<br />
assimilata is identical to M. incrassata and both occur in similar habitats; indeed <strong>the</strong> latter has<br />
<strong>of</strong>ten been considered a variety (var. infuscata) <strong>of</strong> <strong>the</strong> former (e.g. Anderson, 1964; Hertel,<br />
1977). However, M. incrassata differs in several respects, <strong>the</strong> most important <strong>of</strong> which is its quite<br />
different, red-brown hypo<strong>the</strong>cium which lacks any trace <strong>of</strong> purple pigmentation in water, K or<br />
HNO3. In addition, it has an <strong>of</strong>ten darker thallus, less prominent apo<strong>the</strong>cia and a generally<br />
higher proportion <strong>of</strong> septate spores. M. subviolascens is similar in appearance to M. assimilata,<br />
but grows on rock and has a green, K-l- violet epi<strong>the</strong>cial pigment. M. melaenida differs from <strong>the</strong><br />
last three species in its comparatively small apo<strong>the</strong>cia and less conspicuous thallus, absence <strong>of</strong><br />
green pigmentation in apo<strong>the</strong>cial tissues, preponderance <strong>of</strong> 1-septate spores, shorter conidia,<br />
and confinement to argillaceous, or fine-grained, mineral soils. M. crassipes occurs in much <strong>the</strong><br />
same habitats as M. assimilata and <strong>the</strong> two species have been much confused. However, M.<br />
crassipes is easily distinguished by its apo<strong>the</strong>cia which are thinly marginate (at least when young)<br />
and markedly constricted below, <strong>of</strong>ten being turbinate or even short-stipitate; also by its thallus<br />
which is composed <strong>of</strong> small granular-areolae which sometimes proliferate to give it an isidiose<br />
appearance. In section, <strong>the</strong> apo<strong>the</strong>cia <strong>of</strong> M. crassipes display a well-developed excipulum and a<br />
distinctly two-zoned hypo<strong>the</strong>cium.<br />
Pycnidia are rare and difficult to find in members <strong>of</strong> <strong>the</strong> M. assimilata group, and have not yet<br />
been found at all in M. subviolascens. The remaining species appear to have one conidium type<br />
only; although <strong>the</strong> conidia are ra<strong>the</strong>r large, <strong>the</strong>y afe probably microconidia. However, <strong>the</strong><br />
relatively shorter and broader conidia <strong>of</strong> M. crassipes may be better described as mesoconidia.<br />
Cephalodia are here reported from M. assimilata and M. incrassata, apparently for <strong>the</strong> first<br />
time. In both cases <strong>the</strong> blue-green alga concerned is Nostoc, which loses its filamentous form. In<br />
addition, irregular clusters <strong>of</strong> partially disrupted Stigonema filaments are sometimes present<br />
amongst <strong>the</strong> areolae <strong>of</strong> M. assimilata, M. incrassata, and also M. subviolascens. Hyphae,<br />
presumably belonging to <strong>the</strong> lichens, are seen to ramify through <strong>the</strong>se clusters; but whe<strong>the</strong>r or<br />
not <strong>the</strong>se loosely organised structures can be considered to be cephalodia is a problem requiring<br />
fur<strong>the</strong>r detailed anatomical and experimental investigation. Cephalodia or cephalodia-like<br />
structures have not been encountered in any o<strong>the</strong>r Micarea species.<br />
The closely related Lecidea limosa Ach. and L. stenotera (Nyl.) Nyl. have <strong>of</strong>ten been confused<br />
with M. assimilata and M. incrassata, but <strong>the</strong>y can be distinguished by <strong>the</strong>ir pale hypo<strong>the</strong>cia,<br />
paraphyses which are individually supplied with a dense, hyaline, gelatinous sheath, and more<br />
organised excipular structure. Toge<strong>the</strong>r with Catillaria contristans (Nyl.) Zahlbr. <strong>the</strong>y probably<br />
represent a distinct genus within <strong>the</strong> Lecideaceae, although perhaps not far removed from<br />
Micarea. In <strong>British</strong> herbaria (at least) <strong>the</strong>re has been much confusion between M. assimilata and<br />
Lecidea hypnorum Lib. The latter has marginate, brown-black apo<strong>the</strong>cia, simple paraphyses,<br />
0(-3)-septate spores which become straw coloured and finely warted with age, a reddish brown<br />
excipulum composed <strong>of</strong> radiating, stout, pachydermatous, heavily conglutinated hyphae, c.<br />
3-5 /xm wide, and bears little resemblance to a Micarea. In addition, <strong>the</strong> hymenium, hypo<strong>the</strong>cium<br />
and excipulum <strong>of</strong> L. hypnorum (and <strong>the</strong> related L. berengeriana (Massal.) Th. Fr. , and L.<br />
sanguineoatra auct.) <strong>of</strong>ten contain minute granules <strong>of</strong> a dark violet (K-l- aeruginose) pigment.
116 BRIAN JOHN COPPINS<br />
More understandable cases <strong>of</strong> mistaken identity involve Lecidea caesioatra Schaerer, which<br />
bears a close superficial likeness to M. incrassata. It can be distinguished from this Micarea by its<br />
dark blue-grey apo<strong>the</strong>cia which have a caesious bloom when wet, excipulum <strong>of</strong> conglutinated (in<br />
K) hyphae, and non-micareoid phycobiont with cells c. 8-12 /xm diam.<br />
Habitat and distribution: M. assimilata grows on decaying bryophytes and plant detritus on <strong>the</strong><br />
ground or on soil accumulation in rock crevices in arctic (i.e. low altitudes at high latitudes) or<br />
montane situations. The realisation that this species belongs in Micarea came so late in <strong>the</strong><br />
present study, that I have been able to examine only a limited amount <strong>of</strong> herbarium material. To<br />
date, I can confirm its presence in nor<strong>the</strong>rn Scandinavia and <strong>the</strong> single outlying locality in<br />
Scotland. Specimens purporting to be Lecidea assimilata from <strong>the</strong> Alps and o<strong>the</strong>r central<br />
European mountains have proved to be M. crassipes, M. incrassata, or 'Lecidea' species not<br />
referable to Micarea. Material <strong>of</strong> 'Lecidea assimilata'' from Greenland distributed as Hansen<br />
Lich. Greenland Exs. Ill and 247 belongs to Lecidea stenotera and L. limosa respectively.<br />
The Scottish locality is Ben Lawers in Perthshire, a mountain long renowned for its<br />
arctic-alpine flora (Anon, 1972; James, 19656). M. assimilata was collected <strong>the</strong>re by W. L.<br />
Lindsay in 1856. Recent collections from Ben Lawers, and o<strong>the</strong>r Scottish localities, are referable<br />
to o<strong>the</strong>r species, such as Lecidea caesioatra, L. hypnorum, L. limosa, and Micarea incrassata.<br />
Exsiccata: Krypt. Exs. Vindob. 2268 (BM, H). Malme Lich. Suec. 216 (H).<br />
Map 3 Micarea assimilata ® Before 1950 + Micarea incrassata # 1950 onwards
5. Micarea bauschiana (Korber) V. Wirth & Vezda<br />
(Figs5,10A,40D;Map4).<br />
LICHEN GENUS MICAREA IN EUROPE 117<br />
in Vezda & V. Wirth in Folia geobot. phytotax., Praha 11: 95 (1976). _ Biatora bauschiana Korber,<br />
Parerga lich.: 157 (1860). - Lecidea bauschiana (Korber) Lettau in Hedwigia 55: 28 (1914). Type:<br />
Germany, Baden-Wiirttemberg, Baden-Baden, on <strong>the</strong> way to Yburg, on porphyry, 1859, Bausch,<br />
Rabenh. Lich. Eur. 648 (M- lectotype!, sel. V. Wirth & Vezda (loc. cit); M-isolectotype!); additional<br />
isotype material distributed as Arnold Lich. Exs. 120 (BM!, M!).<br />
Lecidea infidula Nyl. in Flora, Jena 51: 475 (1868). Type: Jersey, C. Larbalestier (H-NYL p.m. 5413 -<br />
holotype!).<br />
Lecidea lynceola Th. Fr., Lich. Scand. 2: 561 (1874). Type: Norway, Christiana [Oslo], Tveten, 20 v 1868,<br />
N. G. Moe 257 (UPS -holotype!).<br />
Lecidea semipallens Nyl. in Flora, Jena 59: 234 (1876). Type: Ireland, West Galway, Lough Inagh, 1875, C.<br />
Larbalestier (H-NYL 19399 - lectotype!; isolectotypes: BM ex K!, H-NYL 19402!).<br />
Lecidea dilutiuscula Nyl. in Flora, Jena 59: 308 (1876). Type: England, South Devon, near Buckfastleigh,<br />
H. B. /fo//(BM- lectotype!; H-NYL 10754- isolectotype!).<br />
Lecidea rusticella Nyl. in Flora, Jena 61: 245 (1878). Type: Ireland, West Galway, Connemara, Tullywee<br />
Bridge, 1876, C. Larbalestier (H-NYL 20206- holotype!; topotypes ['1878']: BM!, BM ex K!).<br />
?Lecidea callicarpa Larbal. ex Leighton, Lich. Fl. Br., ed. 3: 266 (1879). Type: Ireland, West Galway,<br />
Glencorbot near Kylemore, 1877, C. Larbalestier (not seen; possibly in BM (incl. BM ex K) but not<br />
traced).<br />
?Lecidea semipallens var. obscurior Lang ex Havaas in Bergens Mus. Arb. 1935(2): 27 (1935). Type:<br />
Norway, Dalsb0, 1903, /. J. Havaas (not seen; not traced in BG or H).<br />
?Catillaria microsporaMas\o\a.in Ukr. bot. Zh. 30(5): 665 (1973). Type: USSR, Regio Volhyniensis, distr.<br />
Ljuboml., in pineto baud procul pag. Kamenca, ad saxa granitica, 11 vi 1969, W. R. Maslova (KWholotype,<br />
not seen; but a slide prepared from <strong>the</strong> holotype by H. Kilias examined).<br />
Thallus thin (
118 BRIAN JOHN COPPINS<br />
<strong>of</strong>ten with a swollen base up to 3 /xm wide; conidia (microconidia) cylindrical, eguttulate,<br />
4-6x0-8-1 fxm. A few collections with pallid apo<strong>the</strong>cia have associated pycnidia, c. 60-100 /xm<br />
diam, with hyaline walls and containing (<strong>of</strong>ten biguttulate) mesoconidia 2-8-4-3X 1-1-3 /xm (see<br />
'Observations' below).<br />
Chemistry: All parts K- , C- , KC— , PD- ; no substances detected by t.l.c.<br />
Observations: Micarea bauschiana is closely related to, and <strong>of</strong>ten confused with, M. sylvicola<br />
(q.v.), and <strong>the</strong> two species <strong>of</strong>ten grow toge<strong>the</strong>r. M. bauschiana is notoriously variable in <strong>the</strong><br />
colour <strong>of</strong> its apo<strong>the</strong>cia, but <strong>the</strong> full range <strong>of</strong> variation from pallid to blackish can be seen over<br />
small distances (a few centimetres) in a single population (Fig. 5: p. 30) and <strong>the</strong> variation appears<br />
to be solely phenotypic.<br />
When growing on iron-rich rocks <strong>the</strong> thallus <strong>of</strong> M. bauschiana is <strong>of</strong>ten oxydated. In a few<br />
specimens (including types <strong>of</strong> Lecidea rusticella) this oxydation has extended to <strong>the</strong> apo<strong>the</strong>cia in<br />
which finely granular, ferruginous material has been deposited (especially in <strong>the</strong> epi<strong>the</strong>cium and<br />
hypo<strong>the</strong>cium); such specimens have been confused with M. lutulata and species <strong>of</strong> Protoblaste-<br />
nia.<br />
A few collections from Austria and Scotland (considered here as M. cf. bauschiana) have<br />
entirely pallid apo<strong>the</strong>cia which are accompanied by pycnidia containing mesoconidia (see<br />
description above). Such pycnidia have not been detected in specimens <strong>of</strong> M. bauschiana with<br />
pigmented apo<strong>the</strong>cia, and it is possible that <strong>the</strong> former collections represent a distinct taxon.<br />
However, one <strong>of</strong> <strong>the</strong>se collections (Austria, Steiemark, Graz, Schockl N <strong>of</strong> St Radegund, 1978,<br />
Poeh (GZU) also has typical (for M. bauschiana) microconidia-containing pycnidia, and it may<br />
be that <strong>the</strong>re is some environmental control (e.g. a response to very low light intensities)<br />
involved in <strong>the</strong> initiation <strong>of</strong> <strong>the</strong> mesoconidial anamorph. Fur<strong>the</strong>r careful field observations and<br />
laboratory studies are required to establish <strong>the</strong> status <strong>of</strong> <strong>the</strong>se seemingly anomalous collections.<br />
Forms <strong>of</strong> M. bauschiana (s. ampl. ) with pallid apo<strong>the</strong>cia should be compared with <strong>the</strong> much<br />
rarer M. lithinella {q. v. ) which can be distinguished by its micareoid phycobiont. In addition, M.<br />
lithinella seems not to occur in <strong>the</strong> ombrophobous, aerohygrophilous Micareetum sylvicolae,<br />
and is a ra<strong>the</strong>r ombrophilous, substratohygrophilous species <strong>of</strong> damp, shaded rocks or stones.<br />
However, I must stress that this ecological interpretation <strong>of</strong> M. lithinella is based on <strong>the</strong> limited<br />
data available from <strong>the</strong> herbarium specimens and requires confirmation by field studies.<br />
Material recently distributed as 'M. bauschiana' in Hertel Lecid. exs. no. 54 is not this species.<br />
It has weakly marginate apo<strong>the</strong>cia due to a well developed excipulum, and a large-celled<br />
(9-17 /xm diam) phycobiont, <strong>the</strong> cells <strong>of</strong> which are deeply penetrated by distinct haustoria (as in<br />
M. intrusa). It is probably an undescribed species <strong>of</strong> Micarea and will be treated in a later<br />
pubhcation following more critical studies.<br />
Habitat and distribution: M. bauschiana is a faithful member <strong>of</strong> <strong>the</strong> Micareetum sylvicolae and<br />
grows on rocks, stones, roots, and consolidated soil in dry underhangs in woodlands or sheltered<br />
valleys. In <strong>the</strong> <strong>British</strong> Isles M. bauschiana is <strong>the</strong> commonest member <strong>of</strong> <strong>the</strong> M. sylvicola group. It<br />
is particularly common in <strong>the</strong> north and west but also occurs in suitable situations (e.g. dry stones<br />
in sandy banks in woodlands) in <strong>the</strong> lowlands <strong>of</strong> south-west England. It seems to be widely<br />
distributed in Europe but I have not seen enough specimens to make an assessment <strong>of</strong> any<br />
distributional tendencies. It occurs in <strong>the</strong> Azores and <strong>the</strong> Canary Islands, but I have not seen any<br />
additional material from outside Europe.<br />
Exsiccata: Arnold Lick. Exs. 120 (BM, M), 1233 (M). Johnson Lick. Herb. AM p.p., 504 (HAMU).<br />
Larbal. Lick. Herb. 68 (BM), 305 p.p. (LD). Rabenh. Lich. Eur. 648 p. max. p. (M). Zwackh Lick.<br />
Exs. 279A-B, 594A-B, 595 (M).<br />
6. Micarea botryoides (Nyl.) Coppins<br />
(Figs 10b, 35, 38A; Map 5)<br />
in D. Hawksw., P. James & Coppins in Lichenologist 12: 107 (1980). - Lecidea apochroeella var.<br />
botryoides Nyl. in Flora, Jena 50: 373 (1867). - Lecidea botryoides (Nyl.) Zahlbr., Ca/. lich. univ. 3: 740
LICHEN GENUS MICAREA IN EUROPE 119<br />
Map 4 Micarea bauschiana # 1950 onwards O Before 1950<br />
(1925). Type: Finland, Tavastia australis, Lammi, Evo, Lapinkallio, 1866, J. P. Norrlin 404 (H-NYL<br />
20685 p.p. -lectotype!; H-isolectotype!).<br />
Note: In <strong>the</strong> protologue Nylander states 'ad lignum putridum' but this is clearly an error (see Vainio<br />
1934: 361). The lectotype has been selected from a packet in <strong>the</strong> Nylander Herbarium containing two<br />
pieces <strong>of</strong> rock - one with M. lutulata, <strong>the</strong> o<strong>the</strong>r (lectotype) with <strong>the</strong> apo<strong>the</strong>cia and characteristic stalked<br />
pycnidia <strong>of</strong> M. botryoides. It is possible that Nylander 's diagnosis was based in part on <strong>the</strong> apo<strong>the</strong>cia <strong>of</strong> M.<br />
lutulata, which bear many similarities to those <strong>of</strong> M. botryoides. Because Art. 70 <strong>of</strong> <strong>the</strong> ICBN is no longer<br />
applicable Lecidea apochroeella var. botryoides cannot be rejected as being based on discordant elements;<br />
consequently <strong>the</strong> part containing M. lutulata is excluded, and that part including stalked pycnidia (as<br />
described in Nylander's diagnosis) is chosen as <strong>the</strong> lectotype.<br />
Thallus effuse, thin, more rarely developing into a thick, loose crust up to c. 0-4 mm thick,<br />
scurfy-granular, pale to dark dull green, dark olive-green, or dull greenish black, sometimes<br />
whitish buff when on dry, deeply shaded rocks. When very thin <strong>the</strong> thallus consists <strong>of</strong> flattened<br />
granules, c. 20-50 ^im diam. , which are <strong>of</strong>ten dispersed and interconnected by white, arachnoid,<br />
prothalline hyphae. Thickening <strong>of</strong> <strong>the</strong> thallus is caused by <strong>the</strong> production <strong>of</strong> goniocysts (c. 16-30<br />
ixm diam), <strong>the</strong> first <strong>of</strong> which appear to arise by budding from <strong>the</strong> primary granules; when thick<br />
<strong>the</strong> thallus has a ± gelatinous appearance when wet. Phycobiont micareoid, cells 4—7 /xm diam.<br />
Apo<strong>the</strong>cia usually few, more commonly absent, immarginate, at first convex-hemispherical,<br />
soon becoming ± globose and much constricted below (sometimes short-stipitate), <strong>of</strong>ten<br />
becoming tuberculate, black, or dark brown (in deep shade), matt, 0- 1-0-25 mm diam, or to 0-5<br />
mm when tuberculate. Hymenium 25-35 ^im tall, hyaline or sometimes tinged dilute brown<br />
(K- , HNO3-) or dilute olivaceous (K- , or K± green intensifying, HNO3+ red) in places, but
120 BRIAN JOHN COPPINS<br />
always with numerous dark brown (K± olivaceous tinge, HN03± reddish tinge) vertical<br />
streaks. Asci cylindrical-clavate or clavate, 20-35x7-9 /xm. Spores ovoid, oblong-ellipsoid or<br />
oblong-ovoid, straight or slightly curved, 0-l(-3)-septate, 8-13(-16)x 2-3-3 •7(-4) /xm. Paraphyses<br />
ra<strong>the</strong>r scanty, <strong>of</strong> two types: p.p. evenly distributed, flexuose, simple or sparingly<br />
branched, sometimes anastomosing, thin, 0-7-1 fxm wide, sometimes widening above to 1 -7 /^m,<br />
walls hyaline throughout; p.p. fasciculate, simple or sparingly branched, stout, c. 2-2-5 /xm<br />
wide, sometimes widening above to 3-5 ixm, coated ± throughout by dark brown pigment.<br />
Hypo<strong>the</strong>cium c. 60-120 /xm tall, dark reddish brown, K- or dulling, HNO3- or red tinge<br />
slightly intensifying; hyphae coated with dark brown pigment, c. 2-3 /xm wide, interwoven but<br />
becoming vertically orientated towards <strong>the</strong> hymenium and sometimes continuing into it as stout,<br />
fasciculate, pigmented paraphyses; ascogenous hyphae similarly pigmented, with short, swollen<br />
cells to 5 /xm wide. Excipulum indistinct, sometimes evident in young apo<strong>the</strong>cia as a reflexed<br />
reddish brown zone concolorous with, or sHghtly paler than, <strong>the</strong> hypo<strong>the</strong>cium; hyphae radiat-<br />
ing, branched and anastomosing, c. 1-1-5 /xm wide, <strong>the</strong>ir walls sometimes with a thin coating <strong>of</strong><br />
brown pigment.<br />
Pycnidia always present and numerous, sessile or, more usually, distinctly stalked, black,<br />
50-400 /xm tall (including stalk) and 40-90 /xm diam; stalks simple, or branched and bearing up<br />
to six pycnidia; <strong>the</strong> 'stalk-part' below <strong>the</strong> current conidia-producing pycnidia <strong>of</strong>ten includes old<br />
pycnidia (see Fig. 35); <strong>the</strong> stalk and pycnidia are usually black but in extreme shade forms <strong>the</strong><br />
stalk tissue may be ± colourless, contrasting with <strong>the</strong> dark brown or blackish, current and old<br />
pycnidia. In microscope preparations (at x400): pycnidiophore tissue dilute to dark fuscous or<br />
reddish brown, K- or dulling, HNO3- or red tinge slightly intensifying; pycnidial wall dark<br />
greenish brown, K- or K-l- green intensifying HN03-(- red. Conidiogenous cells ± cylindrical or<br />
elongate ampuUiform, <strong>of</strong>ten with swollen base which is thickened with brownish pigment, <strong>of</strong>ten<br />
with one or two percurrent proliferations, 3-5-7-5 x 1-1-4 /xm, base sometimes swollen to 2-5 /xm<br />
wide. Conidia (mesoconidia) ± cylindrical, <strong>of</strong>ten biguttulate, sometimes slightly constricted in<br />
<strong>the</strong> middle, 3-5-4-8x1-1-5 /xm.<br />
Chemistry: Thallus K— , C—<br />
detected by t. I.e.<br />
, PD—<br />
; sections <strong>of</strong> apo<strong>the</strong>cia and thallus C— ; no substances<br />
Observations: Micarea botryoides is easily recognised by its numerous, black, stalked<br />
pycnidia, whose walls are dark greenish brown (K- or K-l- green intensifying), and its normal<br />
occurrence on substrate o<strong>the</strong>r than hgnum. It is very similar to <strong>the</strong> rare M. melaeniza, but that<br />
species has shorter conidia, smaller, simple spores, and is apparently confined to lignum. M.<br />
misella <strong>of</strong>ten has stalked, black pycnidia, but is usually hgnicolous. However, it does occasionally<br />
occur on decaying bryophytes where it could be confused with M. botryoides. In such<br />
instances <strong>the</strong>ir separation is easy because <strong>the</strong> pycnidia <strong>of</strong> M. misella contain an olivaceous or dull<br />
brownish pigment that turns violet in K. When on rock <strong>the</strong> apo<strong>the</strong>cia <strong>of</strong> M. botryoides could be<br />
confused with those <strong>of</strong> M. lutulata, but <strong>the</strong> latter species has smaller, simple spores, a<br />
non-micareoid phycobiont, and immersed pycnidia. When on s<strong>of</strong>t lignum it should be compared<br />
with M. nigella which is superficially identical, but has a purple-brown (K-l- green) pigment in its<br />
apo<strong>the</strong>cia and pycnidia.<br />
Sterile forms <strong>of</strong> M. botryoides have puzzled lichenologists for many years, being dismissed<br />
with such remarks as 'indeterminate pycnidia' or 'fungus'. In 1867 Leighton distributed sterile<br />
material <strong>of</strong> M. botryoides in his exsiccate (no. 388), as ' Lecidea sabuletorum var. milliaria (Fr.),<br />
spermagonia', evidently believing it to be <strong>the</strong> pycnidial state oi Micarea lignaria, <strong>the</strong> apo<strong>the</strong>cia<br />
<strong>of</strong> which occur on some <strong>of</strong> his specimens . The<br />
identity <strong>of</strong> <strong>the</strong>se pycnidia remained a mystery until<br />
my discovery <strong>of</strong> fertile material in Scotland in 1976 and my subsequent examination <strong>of</strong> <strong>the</strong> type<br />
material <strong>of</strong> Lecidea apochroeella var. botryoides in 1979.<br />
Habitat and distribution: M. botryoides is usually found as a constituent <strong>of</strong> <strong>the</strong> Micareetum<br />
sylvicolae in dry underhangs, growing on solid rock, loose stones, consolidated soil, tree roots,<br />
and loose mats ot moribund bryophytes; but it also occurs on rock, stones, and decaying<br />
bryophytes in wetter shaded situations. There are a few collections made from <strong>the</strong> s<strong>of</strong>t.
LICHEN GENUS MICAREA IN EUROPE 121<br />
Map 5 Micarea botryoides # 1950 onwards O Before 1950<br />
crumbling lignum <strong>of</strong> old stumps, but I have not seen it on <strong>the</strong> firmer, ± smooth lignum favoured<br />
by M. melaeniza and M. misella. In <strong>the</strong> environs <strong>of</strong> some industrial regions (e.g. <strong>the</strong> West<br />
Yorkshire conurbation) it has been found on bark at <strong>the</strong> bases <strong>of</strong> trunks oi Acer and Betula in<br />
sheltered woodlands.<br />
M. botryoides is a widespread species, especially in western and upland regions <strong>of</strong> Britain, and<br />
is certainly much more common than current records would suggest. Several <strong>of</strong> <strong>the</strong>se records<br />
result from accidental ga<strong>the</strong>rings, being subsequently identified on samples <strong>of</strong> o<strong>the</strong>r members <strong>of</strong><br />
<strong>the</strong> Micareetum sylvicolae. This gives an indication as to how <strong>of</strong>ten this species must have been<br />
overlooked. It is little recorded outside Britain, and to date I know it only from Norway<br />
(Sor-Trondelag and Hordaland), Finland (Tavastia australis), and north-west France.<br />
Exsiccata: Leighton Lich. Brit. 388 (BM, BON, DBN, E, M).<br />
7. Micarea cinerea (Schaerer) Hedl.<br />
(Figs 11, 39; Map 6)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 81, 93 (1892). - Lecidea cinerea Schaerer, Lich.<br />
Heiv. spic, sect. 3: 156 (1826). - Bacidia cinerea (Schaerer) Trevisan in Linnaea 28: 293 (1856). -<br />
Bilimbia cinerea (Schaerer) Korber, Parerga Lich. :<br />
164 (1860). Type: Switzerland, 'ad infimos Abietum<br />
truncos, insylvaKonitz', L. E. Sc/zaerer (G-holotype!; M-? isotype!).<br />
Lecidea sphaeroides var. albella Schaerer, Lich. Helv. spic, sect. 4^5: 165 (1833). Type: Switzerland:<br />
'Schweiz' [on conifer bark], L. E. Schaerer (M - neotype!). See note (i).<br />
Biatora delicatula Korber, Denkschr. Feier ihres funfzigjah. Best, herausg. Schles. Gesellsch. voter. Kult.,
122 BRIAN JOHN COPPINS<br />
Breslau: 233 (1853). - Bilimbia delicatula (Korber) Korber, Syst. lich. Germ: 212 (1855). Type: lectotype<br />
as for Lecidea sphaeroides var. albella Schaerer. See note (ii) below.<br />
Bilimbia cinerea f. hypoleuca Stizenb. ex Arnold in Flora, Jena 58: 598 (1864). - Micarea cinerea f.<br />
hypoleuca (Stizenb. ex Arnold) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 82, 93 (1892).<br />
Type: Germany, Bayern, near Eichstatt, Affenthale, on young Picea abies in a wood, viii 1864, F. C. G.<br />
Arnold (M - lectotype!).<br />
Notes, (i) No material labelled Lecidea sphaeroides var. albella could be traced in G (Geissler, in litt.)<br />
and an undated specimen in M is selected as neotype. If it can be proven that this specimen was collected<br />
before 1833, it should be regarded as a lectotype.<br />
(ii) No material referred to in <strong>the</strong> protologue <strong>of</strong> B. delicatula has been traced in L, WRSL or elsewhere.<br />
However, Korber gives 'Lecidea sphaeroides a albella Schaer. Enum. 193.' as a synonym. This citation<br />
refers to Schaerer (1850) and is an indirect reference to Lecidea sphaeroides var. albella Schaerer (1833).<br />
Consequently I have typified both names with <strong>the</strong> same specimen in M.<br />
Thallus effuse, sometimes partly immersed in <strong>the</strong> substratum (especially when on lignum),<br />
more usually developed on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> substratum as rounded, shallow-convex, hemispherical<br />
or ± globose areolae. Areolae scattered to ± contiguous, smooth, greenish white to<br />
blue-grey, or more rarely becoming dark grey, <strong>of</strong>ten dark coloured on upper surface but<br />
greenish white below, c. 40-160 fxva diam or up to 300 jxm diam if containing a pycnidium <strong>of</strong> <strong>the</strong><br />
macroconidial anamorph. Areolae in section, ecorticate but with a hyaline amorphous covering<br />
layer c. 2-5 /xm thick; outermost hyphae <strong>of</strong>ten with grey-green to aeruginose walls, K-,<br />
HNO3-I- red. Phycobiont micareoid, cells 4-7 /xm diam.<br />
Apo<strong>the</strong>cia usually numerous (see 'observations'), adnate, plane to convex, sometimes<br />
becoming tuberculate, sometimes with an indistinct margin that is ± flush with <strong>the</strong> level <strong>of</strong> <strong>the</strong><br />
disc, pale leaden-grey to grey-black (margin <strong>of</strong>ten paler), or ivory-white or pallid in shade forms,<br />
(0-2)0-3-0-7 mm diam or up to 1-3 mm when tuberculate. Disc matt and finely roughened, but<br />
margin smooth and <strong>of</strong>ten ± glossy. Hymenium 55-70 /xm tall, hyaline, but usually olivaceous or<br />
aeruginose (K— , HNO3+ red) in upper part (epi<strong>the</strong>cium). Asci clavate 50-65x15-20 /am.<br />
Spores fusiform, <strong>of</strong>ten slightly curved, (3-)5-7-septate, (19-)23-34(-38)x 4-5-6 ixm. Paraphyses<br />
numerous, branched and <strong>of</strong>ten anastomosing, 1-1-4 /xm wide; apices <strong>of</strong>ten more richly<br />
branched and entangled, <strong>of</strong>ten slightly incrassate to c. 1-8 ^im, or 2-5 /xm due to thickening by<br />
greenish pigment. Hypo<strong>the</strong>cium c. 40-70 /xm tall, hyaline; hyphae interwoven, c. 2-4 /xm wide.<br />
Excipulum well developed, hyaline or pale straw, or richly branched and anastomosing hyphae,<br />
c. 1-1-5 /xm.<br />
Pycnidia frequently present, <strong>of</strong> two types: (a) immersed in areolae, white or faintly greenish<br />
around <strong>the</strong> ostiole, 160-300 /xm diam, ostiole <strong>of</strong>ten widely gaping; conidia (macroconidia) ±<br />
straight or flexuose, filiform, 9-17-septate, 50-1 10 xc. 1 /xm; (b) ± sessile, white, c. 40-70 /xm<br />
diam, ostioles not, or only slightly, gaping; conidia (microconidia) narrowly fusiform-cylindrical<br />
(3-8-)^5xO-5-0-7/xm.<br />
Chemistry: Thallus and whitish apo<strong>the</strong>cia K-, C-l- red, PD-; apo<strong>the</strong>cia in section C-l- red;<br />
t.l.c: gyrophoric acid.<br />
Observations: M. cinerea is very variable with regard to <strong>the</strong> colour <strong>of</strong> its apo<strong>the</strong>cia and thallus;<br />
as with M. peliocarpa this variation is due to <strong>the</strong> amount <strong>of</strong> green pigment produced in response<br />
to environmental factors, especially light. Morphologically and chemically M. cinerea is closely<br />
allied to M. alabastrites and M. peliocarpa; <strong>the</strong> diagnostic features <strong>of</strong> each <strong>of</strong> <strong>the</strong>se three species<br />
are compared in Table 4. Forms <strong>of</strong> M. cinerea with blackish apo<strong>the</strong>cia can be confused with M.<br />
lignaria (including var. endoleuca) but on close inspection <strong>the</strong> latter will be seen to differ in its ±<br />
globose apo<strong>the</strong>cium with a poorly differentiated excipulum (in vertical section), usually<br />
brownish or greenish upper hypo<strong>the</strong>cium, less strongly branched paraphyses and chemistry.<br />
M. cinerea is usually well fertile with numerous apo<strong>the</strong>cia, but sterile forms with numerous<br />
macroconidia-containing pycnidia have been encountered; <strong>the</strong> macroconidia are about twice as<br />
long as those found in M. peliocarpa and M. alabastrites and so are readily identifiable (see 'Key<br />
to species without apo<strong>the</strong>cia, pp. 107-108).
LICHEN GENUS MICAREA IN EUROPE 123<br />
Habitat and distribution: In <strong>the</strong> <strong>British</strong> Isles M. cinerea is mostly found in communities <strong>of</strong> (or<br />
closely akin to) <strong>the</strong> Parmelietum laevigatae on <strong>the</strong> trunks or over bryophytes <strong>the</strong>reon <strong>of</strong> Quercus,<br />
Betula and, less <strong>of</strong>ten, Alnus, Corylus, Fraxinus, Ilex, Sorbus, Larix, Pinus, and Pseudotsuga.<br />
Associated species in <strong>the</strong>se communities include Catillaria pulverea, Cladonia coniocraea, C.<br />
macilenta, C. squamosa, Haematomma caesium, H. elatinum, Lecidea icmalea, Lepraria incana,<br />
Micarea alabastrites, M. stipitata, M. syno<strong>the</strong>oides, Mycoblastus sterilis, Parmelia laevigata, P.<br />
saxatilis, Platismatia glauca, Stenocybe septata, Thelotrema lepadinum, Trapelia corticola ined. ,<br />
Frullania tamarisci, Lejeunea ulicina, Scapania gracilis, Dicranum fuscescens, and Hypnum<br />
cupressiforme. Less commonly it is found in <strong>the</strong> Ugnum <strong>of</strong> fallen, decorticated trunks. Occurrences<br />
on o<strong>the</strong>r substrata are rare although I have found it on one occasion growing directly on<br />
rock (Coed Hafod in Denbigh) where it was associated with Trapelia involuta on <strong>the</strong> top <strong>of</strong> a<br />
dry-stone wall in an oak-birch wood. In addition, it was found on mosses on epidiorite at an<br />
altitude <strong>of</strong> 1000 m in <strong>the</strong> Ben Alder range <strong>of</strong> Inverness-shire by Dr O. L. Gilbert; <strong>the</strong> specimen is<br />
sterile but has pycnidia with <strong>the</strong> characteristic macroconidia. In Britain M. cinerea usually occurs<br />
at lower altitudes (mostly below 300 m), and exhibits a distribution pattern attributable to <strong>the</strong><br />
General Western Group <strong>of</strong> Coppins (1976). However, <strong>the</strong>re are two outlying easterly localities<br />
(in North Northumberland and East Perth) both <strong>of</strong> which are sheltered, more or less undisturbed,<br />
valley woodlands.<br />
From outside Britain I have seen material <strong>of</strong> M. cinerea from Hordaland and Rogaland in<br />
western Norway, <strong>the</strong> east Sudety and Vysoke Tatry <strong>of</strong> Czechoslovakia, sou<strong>the</strong>rn Germany, and<br />
<strong>the</strong> Swiss, Austrian and Italian Alps. Most collections from sou<strong>the</strong>rn Germany and <strong>the</strong> Alps are<br />
Map 6 Micarea cinerea • 1950 onwards O Before 1950
124 BRIAN JOHN COPPINS<br />
from <strong>the</strong> trunks <strong>of</strong> conifers (Picea, Pinus, and Abies), but <strong>the</strong>y also included specimens from <strong>the</strong><br />
trunks <strong>of</strong> A Inus and Betula, lignum <strong>of</strong> stumps, fallen trees, and old fence posts. In addition, it<br />
was collected several times on thin twigs <strong>of</strong> Picea by F. Arnold and his contemporaries, and it is<br />
possibly <strong>of</strong> interest to note that I do not know it as an inhabitant <strong>of</strong> twigs in north-west Europe.<br />
Unlike M. alabastrites and M. peliocarpa, M. cinerea is not known from Macaronesia, and I<br />
have not seen any material <strong>of</strong> it from outside Europe.<br />
Exsiccata: Arnold Lich. Exs. 548 (M), 549 (BM ex K, M). Arnold Lich. Mon. 47 (BM ex K), 1 15 (BM ex<br />
K, M, MANCH), 116 (BM ex K, M). Britz. Lich. Exs. 846 (M). Hepp Flecht. Eur. 21 p.p. (BM, M). Lojka<br />
Lich. Hung. 60 (M). Vezda Lich. Set. 1087 (BM, S). Zwackh Lich. Exs. 898 (M).<br />
8. MicareacontextaHedl.<br />
(Figs 12A, 40A-B)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 83, 96 (1892). - Catillaria contexta (Hedl.) Zahlbr,<br />
Cat. lich. univ. 4: 35 (1926). Type: Sweden, Halsingland, Ovanaker, 1891, J. T. Hedlund (S -<br />
lectotype!).<br />
Thallus effuse, endoxylic, inapparent or evident as a slight bleaching <strong>of</strong> <strong>the</strong> wood, consisting<br />
<strong>of</strong> scattered, small, rounded granules (c. 15-25 pm diam) buried between <strong>the</strong> wood fibres;<br />
external hyphae <strong>of</strong> granules with dark green walls, K— , HNO3-I- red. Phycobiont micareoid,<br />
cells 4-7 pm diam.<br />
Apo<strong>the</strong>cia numerous, immarginate, ± globose from <strong>the</strong> start, <strong>of</strong>ten becoming tuberculate,<br />
black, matt, 0- 1-0-2 mm diam, or to 0-3 mm diam when tuberculate. Hymenium 35-45 pm tall;<br />
upper part (epi<strong>the</strong>cium) dark green, K- or green intensifying, HNO3+ purple-red; remaining<br />
(lower) part ± hyaline or dilute greenish with dark green, vertical streaks, sometimes with a few,<br />
minute, purple-violet (K-l- aeruginose) granules. Asci clavate, 35-40x10-14 pm. Spores ovoid<br />
or oblong-ovoid, upper cell broader than <strong>the</strong> lower and with a more rounded apex, 1-septate, or<br />
rarely with a thin, additional septum in <strong>the</strong> lower cell, 7-13(-14)x(2-3-)3-4-5 pm. Paraphyses<br />
ra<strong>the</strong>r scanty, <strong>of</strong> two types: p. max. p. branched and anastomosing, haline, thin, 0-6-0-8 pm<br />
wide, apices <strong>of</strong>ten with thickened, pigmented walls and <strong>the</strong>n to 2 pm wide; /?. min. p. scattered<br />
or in small fascicles, simple, with thickened, pigmented walls throughout and 1-5-2 pm wide,<br />
apices sometimes widening to 3 pm. Many apices overtopping <strong>the</strong> asci to form a ± distinct<br />
epi<strong>the</strong>cium. Hypo<strong>the</strong>cium 20-90 pm tall, dark green or dark purple (colours <strong>of</strong>ten intermixed),<br />
K-l- dark olive-green or aeruginose, HNO3-I- purple-red; hyphae interwoven, hyaline or<br />
thickened with pigment, 1-2 pm wide; ascogenous hyphae with swollen cells to 4 pm wide.<br />
Excipulum indistinct, sometimes evident as a narrow, dark greenish zone, c. 5-12 /am wide,<br />
forming a lateral border to <strong>the</strong> reflexed edge <strong>of</strong> <strong>the</strong> hymenium.<br />
Pycnidia usually present but inconspicuous, immersed between <strong>the</strong> wood fibres or emergent<br />
to sessile, black, with dark greenish walls, K - or green intensifying, HNO3-I- purple-red; <strong>of</strong><br />
two types (a) c. 40 pm diam; conidia (mesoconidia) cylindrical, <strong>of</strong>ten faintly biguttulate<br />
3-8^-7x 1-3-1-8 pm; (b) c. 20-40 pm diam; conidia (microconidia) narrowly cylindrical, 4-5x<br />
0-8-1 pm.<br />
Chemistry: Apo<strong>the</strong>cia sections C— ; material insufficient for analysis by t.l.c.<br />
Observations: Micarea contexta is characterised by its endoxylic thallus, very small, black, ±<br />
globose apo<strong>the</strong>cia, dark green epi<strong>the</strong>cium, dark greenish or purplish hypo<strong>the</strong>cium, ovoid,<br />
1-septate spores, thin hyaline paraphyses and an absence <strong>of</strong> stalked pycnidia. It is apt to be<br />
confused with diminutive, immature forms <strong>of</strong> M. melaena, but that species has more numerous<br />
paraphyses, longer microconidia, and a superficial, granular thallus which contains gyrophoric<br />
acid when in a healthy condition. When mature, M. melaena has more robust apo<strong>the</strong>cia and<br />
larger, 3-septate spores. M. eximia can be distinguished from M. contexta by its more brightly<br />
coloured epi<strong>the</strong>cium and narrower, ± fusiform spores; and M. nigella can be distinguished by its<br />
simple spores and stalked pycnidia. M. olivacea differs in having more numerous paraphyses<br />
which are broader (1-1-2 ^im) when hyaline, a complete absence <strong>of</strong> purple pigmentation in its<br />
apo<strong>the</strong>cia, and relatively narrower, ± oblong spores.<br />
1
LICHEN GENUS MICAREA IN EUROPE 125<br />
Habitat and distribution: M. contexta occurs on conifer lignum, probably in sheltered,<br />
woodland situations. Species associated with M. contexta on <strong>the</strong> specimens examined include<br />
Arthonia helvola, Cetraria pinastri, Cladonia spp., (scattered squamules), Hypogymnia phy-<br />
sodes, Lecanora symmicta agg., Lecidea efflorescens, L. pullata, Micarea anterior, M. misella,<br />
Parmeliopsis ambigua, and P. hyperopta.<br />
The known distribution <strong>of</strong> M. contexta is restricted to middle Sweden, from where it was<br />
found in several localities by Hedlund. Supposed collections from outside Sweden seen by me<br />
belong to M. melaena. Never<strong>the</strong>less, M. contexta should be sought for in o<strong>the</strong>r regions,<br />
especially those containing naturally occurring coniferous forests.<br />
Exsiccata: Malme Lich. Suec. 28 (M, S).<br />
9. Micarea crassipes (Th. Fr.) Coppins, comb. nov.<br />
(Figs4B, 12B,41C)<br />
Helocarpon crassipes Th. Fr., Lich. arctoi: 178 (1860); and in Nova Acta R. Soc. Sclent. Upsal. Ill, 3: 278<br />
(1861). - Lecidea crassipes (Th. Fr.) Nyl. in Flora, Jena 45: 464 (1861). Type: Norway, Finnmark,<br />
Aldjok, 1857, Th. M. Fries (S-lectotype!; isolectotypes: BM!, BM ex K!, LD, UPS!).<br />
Lecidea crassipes f. morlformls Th. Fr., Lich. Scand. 2: 520 (1874). Type: Norway, Finnmark, Masoy, 16<br />
vii 1864, Th. M. Fries (UPS-lectotype!).<br />
Lecidea crassipes f. pulverula Th. Fr., Lich Scand. 2: 520 (1874). Type: Norway, Sor-Trondelag, Oppdal<br />
hd., Dovre, Kongsvoll, Hogsnyta, 11 viii 1863, Th. M. Fries (UPS-lectotype!).<br />
Lecidea hypopodla f. subasslmllata Nyl. in Bull. Soc. linn. Normandle, IV, 1: 242 (1887). Type: U.S.A.,<br />
Alaska, Bering Strait, St Lawrence Island, 1879, E. Almqulst (H-NYL 20915 - holotype!).<br />
Thallus growing on bryophytes or plant debris, rarely spreading on to sandy soil, composed <strong>of</strong><br />
small, ± globose areolae. Areolae cream-white, pale grey-brown, sometimes ash-grey in part,<br />
matt, ra<strong>the</strong>r fragile, 0-07-0-2 mm diam; <strong>the</strong> larger areolae sometimes budding to produce<br />
smaller, secondary granules <strong>the</strong>reby giving <strong>the</strong> thallus an isidiose appearance. Areolae in section<br />
without a hyaline covering layer; thallus hyphae entirely hyaline, c. 2-3 pm wide. Phycobiont<br />
micareoid, cells 4-7 pm diam.<br />
Apo<strong>the</strong>cia numerous, sessile but (in section) markedly constricted below, or turbinate, or<br />
short-stalked, 0-2-0-6 mm diam; disc matt, at first plane with a thin, <strong>of</strong>ten slightly raised,<br />
concolorous or slightly paler, <strong>of</strong>ten ± glossy, margin, 0-02-0-05 mm wide; disc later expanding,<br />
becoming convex and excluding <strong>the</strong> margin. On rare occasions apo<strong>the</strong>cia become tuberculate<br />
and up to 1-4 pm diam. Hymenium 45-50 pm tall; upper part (epi<strong>the</strong>cium) dark green or<br />
olivaceous and K-h green-intensifying, or purple-brown and K+ purple-intensifying; remaining<br />
(lower) part dilute purplish, K+ purple-intensifying or K+ sordid-green. Asci clavate, 40-<br />
48x11-14 pm. Spores 0(-l)-septate, fusiform-ellipsoid, (9-)10-17(-21)x(2-5-)3-4-5 pm.<br />
Paraphyses numerous, simple, or sparingly branched above, sometimes anastomosing, 1-1-5<br />
ju-m wide, sometimes widening above to 2-5 ^tm; apical walls hyaline but <strong>of</strong>ten surrounded by<br />
dense pigment. Hypo<strong>the</strong>cium dark, variably pigmented: upper part (c. 30-60 pm) dark<br />
purple-brown, K-l- dark olive-green, or K-l- purple-intensifying in part or in whole; lower part<br />
(including 'stipe') paler and mottled, brown- or vinose-red and K- , or with purple tinge and K+<br />
purple-intensifying; hyphae in upper part tightly interwoven but becoming vertically orientated<br />
towards <strong>the</strong> hymenium, c. 1-5-2 pm wide, embedded in a densely pigmented gel-matrix,<br />
intermixed with ascogenous hyphae c. 2-5-4 ^tm wide; hyphae in lower part loosely interwoven,<br />
becoming outwardly directed laterally (i.e. as <strong>the</strong>y approach <strong>the</strong> excipulum), c. 1-1-5 pm wide,<br />
embedded in a gel-matrix that swells and partially dissolves in K. Excipulum well developed,<br />
and conspicuous in young mature apo<strong>the</strong>cia, c. 50 pm wide laterally; innerpart concolorous with<br />
upper hypo<strong>the</strong>cium; outer part dilute to dark olivaceous, K+ green-intensifying; hyphae<br />
radiating, branched and anastomosing, c. 1-1-5 pm wide. Excipular and hypo<strong>the</strong>cial tissues<br />
usually extended vertically downwards to form a 'stipe' (Fig. 4B).<br />
Pycnidia rare, sessile, black, c. 60-150 pm diam, ostiole sometimes gaping; walls c. 10-<br />
17 pm wide, green-black (K- or K+ green-intensifying) in upper parts, changing to dark<br />
red-brown (K-) below; conidiogenous cells elongate-ampulliform, strongly tapered towards
126 BRIAN JOHN COPPINS<br />
<strong>the</strong> neck, 6-9x3-4 /xm, neck 1-1-3 /xm wide; conidia {7 mesoconidia) cylindrical, 4-5-5-7xl-2-<br />
1-5 ixm.<br />
Chemistry: Thallus K— , C-, KC-, PD-; t.l.c: no substances.<br />
Observations: The combination <strong>of</strong> a finely granular (sometimes isidiose) whitish to grey,<br />
muscicolous thallus and turbinate or short-stalked, marginate (when young), black apo<strong>the</strong>cia<br />
make this an easy species to recognise, even with a hand lens. Internally, <strong>the</strong> apo<strong>the</strong>cia have a<br />
well differentiated excipulum and a distinctly two-zoned hypo<strong>the</strong>cium. This structure, which is<br />
ra<strong>the</strong>r 'advanced' for a Micarea, has led me to deliberate on <strong>the</strong> merits <strong>of</strong> <strong>the</strong> monotypic genus<br />
Helocarpon Th. Fr. However, characters such as <strong>the</strong> basic arrangement, organisation and form<br />
<strong>of</strong> apo<strong>the</strong>cial hyphae (including paraphyses), ascus structure, morphology <strong>of</strong> pycnidia, and<br />
thallus structure (including phycobiont), are still those <strong>of</strong> a Micarea. The vertical extension <strong>of</strong><br />
excipular and hypo<strong>the</strong>cial tissues to produce a 'stipe' is not unique to M. crassipes, and is found<br />
in some specimens <strong>of</strong> M. lignaria and M. ternaria, and in an undescribed species from New<br />
Zealand (North Island: Trounson Kauri Reserve, on hepatics on rotten stumps 1967, D. M.<br />
Henderson (E)). This last species closely resembles M. crassipes (e.g. apo<strong>the</strong>cia marginate when<br />
young, apo<strong>the</strong>cial pigmentation, thallus type) but has broader, ellipsoid or ovoid, 1-septate<br />
spores (13-16-5x5-5-7 /Am) and ra<strong>the</strong>r sparse paraphyses.<br />
Overall considerations <strong>of</strong> morphology and ecology lead me to assign M. crassipes to <strong>the</strong> group<br />
including M. assimilata, and it is to <strong>the</strong> account <strong>of</strong> that species <strong>the</strong> reader is referred for fur<strong>the</strong>r<br />
discussions.<br />
Habitat and distribution: M. crassipes grows over bryophytes and plant debris on <strong>the</strong> ground<br />
and amongst rocks in nor<strong>the</strong>rn Fennoscandia and at high altitudes (probably above 1000 m) in<br />
<strong>the</strong> Alps and o<strong>the</strong>r central European mountains. From Norway and Sweden I have seen several<br />
collections from north <strong>of</strong> c. 63°N and in Russia it occurs on <strong>the</strong> Kola Peninsula. In Finland it<br />
extends to <strong>the</strong> sou<strong>the</strong>rnmost parts (Karelia australis, c. 60°30'N). In central Europe it occurs in<br />
<strong>the</strong> western Sudety (Krkonose, at 1000 m) <strong>of</strong> Czechoslavakia, and southwards its range extends<br />
east to <strong>the</strong> Alps (Austrian Tirol, and Dolomiti in north Italy) and west to <strong>the</strong> Transylvanian Alps<br />
(Munjii Retezat, at c. 2000-2500 m) <strong>of</strong> Romania. From outside Europe it has been collected on<br />
St Lawrence Island in <strong>the</strong> Bering Strait; it is quite likely to have a circumpolar distribution.<br />
Exsiccata: Arnold Lich. Exs. 556 [A] (BM ex K, H, H-NYL 16578), 556B (BM ex K, H, H-NYL p.m.<br />
5105), 1121 (BM ex K). Fellman Lich. Arct. 165 (BM ex K, H, H-NYL 16570). Malme Lich. Suec. 362<br />
(BM). NorrlinandNyl. Herb. Lich. Fenn. 194A, B(H). Vezda L/c/i. Bohem. 282(LD). Vezda L/c/i. Sel. 11<br />
(BM).<br />
10. Micarea curvata Coppins, sp.nov.<br />
(Fig. 12C)<br />
Thallus probabiliter albido-griseus vel brunneo-griseus, granuloso-verruculosus. Algae cellulis 4-7 /Am<br />
diam. Apo<strong>the</strong>cia immarginata, convexo-hemisphaerica mox tuberculata, atrobrunnea, 0-2-0-5 mm diam,<br />
aut ad 0-65 diam ubi tuberculata. Hymenium c. 60 /tm altum, ± hyalinum, cum vittis verticalibus, pallide<br />
fuscis; parte summa (epi<strong>the</strong>cio) pallide fuscis, K-, Ascosporae fabiformes vel valde curvatae,<br />
(O-)l-septatae, 9-ll-7x2-5-3-8 /^m. Paraphyses ramosae et anastomosantes, graciles, c. 0-8-1 /xm latae,<br />
apicibus baud incrassatis et baud pigmentiferis. Hypo<strong>the</strong>cium pallidum. Excipulum reflexum, pallidum,<br />
margine externo pallide fusescenti. Pycnidia ignota. Thallus et apo<strong>the</strong>cia in sectione C+ rubra.<br />
Typus: Germania, Guestphalia, Gravenhorst, in muro lapideo ad lapidem arenarium, leg. Th. R. J.<br />
Nitschke (WRSL -holotype )<br />
!<br />
.<br />
Thallus probably grey or brownish grey and granular-verrucose, but difficult to interpret in<br />
<strong>the</strong> single specimen in which <strong>the</strong> thallus is invaded by foreign algae and dematiaceous hyphae.<br />
Phycobiont micareoid, cells 4-7 pm diam.<br />
Apo<strong>the</strong>cia convex-hemispherical to tuberculate, immarginate, dark brown, 0-2-0-5 mm diam,<br />
or to 0-65 mm diam when tuberculate. Hymenium c. 60 pm tall ± hyaline with pale fuscous<br />
brown vertical streaks; upper part (epi<strong>the</strong>cium) pale fuscous brown, K- , HNO3. Asci clavate, c.<br />
45-50x10-13 pm. Spores fabiform or distinctly curved, (0-) 1-septate, 9-11-7x2-5-3-8 pm.
LICHEN GENUS MICAREA IN EUROPE 127<br />
Paraphyses branched and anastomosing c. 0-8-1 /xm wide, not swollen or pigmented at apices.<br />
Hypo<strong>the</strong>cium pale, tinged dilute straw-brown. Excipulum soon reflexed, internally pale with<br />
straw-brown tinge, becoming slightly darker (pale fuscous) towards <strong>the</strong> outer edge; hyphae<br />
radiating, branched and anastomosing, c. 0-8-1 /xm wide.<br />
Pycnidia not found.<br />
Chemistry: Sections <strong>of</strong> thallus C-l- red, PD-; sections <strong>of</strong> apo<strong>the</strong>cia C-l- red; probably<br />
containing gyrophoric acid but material insufficient for analysis by t.l.c.<br />
Observations: M. curvata is characterised by its distinctly curved, 1-septate spores, fuscous<br />
brown pigment in <strong>the</strong> apo<strong>the</strong>cia which is unchanged by K or HNO3, and C+ red reactions<br />
(? gyrophoric acid) <strong>of</strong> apo<strong>the</strong>cia and thallus in sections. It is reminiscent <strong>of</strong> M. subnigrata but<br />
that species has uncurved, ellipsoid spores and all parts C-. In outward appearance M. curvata<br />
is similar to some forms <strong>of</strong> <strong>the</strong> almost ubiquitous Scoliciosporum umbrinum, with which it could<br />
be mistaken in <strong>the</strong> field.<br />
Habitat and distribution: M. curvata is known only from a single collection on sandstone in <strong>the</strong><br />
Wroclaw Herbarium, which is labelled '?Biatora nov. sp. Auf einer Steinmauer bei<br />
Gravenhorst/Westfalen leg. Nitschke' (label kindly transliterated by Pr<strong>of</strong>. Hertel). No recognisable<br />
associate species are present on <strong>the</strong> small specimen although <strong>the</strong>re are a few, rounded,<br />
white areolae (C+ red) probably belonging to Trapelia coarctatas. ampl. From <strong>the</strong> appearance<br />
<strong>of</strong> <strong>the</strong> specimen it occurred in a humid, sheltered, possibly shaded or north-facing situation.<br />
11. Micarea denigrata (Fr.) Hedl.<br />
(Figs IB, 13, 42; Map 7)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 89 (1892). - Biatora denigrata Fr. in K. svenska<br />
VetenskAkad. Handl. 1822: 265 (1822). - Catillaria denigrata (Fr.) Boistel, Nouv. Fl. Lich. 2: 199 (1903).<br />
Type: Sweden, Smaland, 'Femsjo in parietibus vetustus', E. M. Fries (UPS - lectotype! [t.l.c:<br />
gyrophoric acid]).<br />
Lecidea anomala f. pyrenothizans Nyl., Lich. Scand.: 203 (1861). - Micarea denigrata f. pyrenothizans<br />
(Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 89 (1892). Type: Finland, Nylandia,<br />
Helsingfors [Helsinki], on lignum, 1860, W. A^>'/flnrfer(H-NYL 21665 -lectotype!).<br />
Lecidea parissima Nyl. in Crombie mJ. Bot. , Lond. 9: 178 (1871), and in/. Linn. Soc. Bot. 11: 484 (1871).<br />
Type: England, Middlesex, Hendon, on old pales, 1870, J. M. Crombie (H-NYL 21659 - holotype!;<br />
isotypes: BM!, BM ex K!).<br />
Lecidea spodiza Nyl. in Flora, Jena SI: 9 (1874). Type: Scotland, Perthshire, near Killin, on old worked<br />
conifer lignum, 1873,7. M. Crom6/e (H-NYL 21734- lectotype!; BM-isolectotypes!).<br />
Lecidea hemipoliella Nyl. in Flora, Jena 58: 11 (1875). - Micarea hemipoliella (Nyl.) Vezda in Vezda & V.<br />
Wirth in Folia geobot. phytotax., Praha 11: 100 (1976). - Micarea denigrata f. hemipoliella (Nyl.) Hedl.<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3) 78, 89 (1892). Type: Finland, Tavastia australis, Evo,<br />
± smooth bark oiAlnus, 1873, /. P. Norrlin (H - lectotype!, isolectotypes: H(x3)!, H-NYL 21691!).<br />
Lecidea discretula Nyl. in Flora, Jena 58: 444 (1875). Type: Romania, 'Circa Thermas Herculis in Banatu<br />
(Hung.), supra Daedaleam in trunco quercino', 1874, H. Lojka (H-NYL 21693 - holotype!).<br />
Lecidea aniptiza Stirton in Rep. Trans. Glasgow Fid Nat. 4: 85 (1876). Type: Scotland, Perthshire,<br />
Killiecrankie, on lignum, 1875, /. Stirton (GLAM - lectotype!; BM - isolectotype!).<br />
Lecidea denigrata var. submisella Nyl. in Vainio in Medd. Soc. Fauna Fl. fenn. 3: 112 (1878). - Catillaria<br />
syno<strong>the</strong>a f. submisella (Nyl.) Blomb. & Forss., Enum. PI. Scand.: 92 (1880). Type: Finland, Tavastia<br />
australis, Korpilahti, on lignum, 1874, E. A. Lang [Vainio] (H-NYL 21661 - lectotype!).<br />
Lecidea spodiza f. ecrustacea Lamy in Bull. Soc. bot. Fr. 25: 440 (1878). Type: France, Puy-de-D6me,<br />
Monte Dore, near Cascade du Queureilh, on lignum <strong>of</strong> conifer trunk, 31 vii 1878, Lamy (H-NYL 21731 -<br />
lectotype!).<br />
Lecidea praeviridans Nyl. , Suppl. Lich. env. Paris: 5 (1879). - Biatorina praeviridans (Nyl.) Boistel, Nouv.<br />
Fl. Lich. 2: 194 (1903). - Catillaria praeviridans (Nyl.) Zahlbr., Cat. Lich. univ. 4: 64 (1926). Type:<br />
France, Haute Loire, Saugues, on Pinus bark, 18 -, A. Boistel (H-NYL 19221 - lectotype! [t.l.c:<br />
gyrophoric acid]; BM - isolectotype!).<br />
Lecidea denigrata f. sublivescens Nyl. in Flora, Jena 64: 539 (1881). Type: Romania, Maramures, 'prope
128 BRIAN JOHN COPPINS<br />
balneum kabola Pojana, com. Marmaros in Hung.', on bark <strong>of</strong> Pinus sytvestris, H. Lojka 4455, 'Lich.<br />
Hung. exs. (ined.) n. 305. ad int.' (H-NYL 11663 -holotype!).<br />
Lecidea denigrata f. pseudoglomerella Harm, in Bull. Seanc. Soc. Sci. Nancy. II, 33: 58 (1899 ['1898']). -<br />
Catillaria denigrata var. pseudoglomerella (Harm.) Boistel, Nouv. Fl. Lich. 2: 199 (1903). Type: France,<br />
Meur<strong>the</strong>-et-Moselle, La Malgrange, on oak posts [lignum], 8 v 1894, /. Harmand, Lich, Loth. 838p.p.<br />
(ANGUC-lectotype! [t.l.c: gyrophoricacid]; DUKE-isolectotype!).<br />
Catillaria prasina f. /o«g/or Erichsen in Schr. naturw. Ver. Schleswig-Holst. 11: 101 (1937). Type: West<br />
Germany, Schleswig-Holstein, Lauenberg, Sachsenwald, Rev. Kl. Viert, on roots <strong>of</strong> old Fagus, 2 xii<br />
1934, C. F. E. Erichsen (HBO -holotype!).<br />
Micarea denigrata var. friesiana Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 89 (1892);<br />
nam. inval. (Art. 26).<br />
Micarea denigrata war . friesiana f. vulgaris Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 79, 90<br />
(1892); nom. inval. (Art. 26).<br />
Micarea andesitica Vezda in Poelt & Vezda, Bestimmungsschl. europ. Flechten. Erganzungsheft I: 160<br />
(1977); nom. nudum (Art. 32). Spec, orig.: Hungary, Matra, Matrafiired, Pipis-Legy, 380 m, on<br />
andesitic rock, 17 xi 1976, A. Kiszely & A. Vezda (hb Vezda!).<br />
? Lecidea hemipoliella* semialbula Stirton in Rep. Trans. Glasgow Fid Nats. 4: 89 (1876). - Biatorina<br />
syno<strong>the</strong>a var. semialbula (Stirton) A. L. Sm., Monogr. Br. Lich. 2: 122 (1911). Type: Scotland,<br />
Su<strong>the</strong>rland, Altnaharra, on hgnum, Stirton (not seen; not traced in BM or GLAM).<br />
?Catillaria syno<strong>the</strong>a f. major B. de Lesd., Rech. Lich. Dunkerque, Suppl. 1: 118 (1914). Type: France,<br />
Nord, Dunkerque, Ghyvelde, on piece <strong>of</strong> wood (Pinus) on <strong>the</strong> sand in a dune, B. de Lesdain (not seen).<br />
?Catillaria syno<strong>the</strong>a f. fusca B. de Lesd., Rech. Lich. Dunkerque, Suppl. 1: 118 (1914). Type: France,<br />
Nord, Dunkerque, St-Pol, on a stake, B. de Lesdain (not seen).<br />
Lecidea syno<strong>the</strong>a auct., non Ach. (1808). See note below.<br />
Note: Lecidea syno<strong>the</strong>a Ach. in K. svenska VetenskAkad. Handl. 1808: 236 (1808). From <strong>the</strong> comments<br />
on Acharian specimens by Hedlund (1892: 91) and my examination <strong>of</strong> Acharian material in BM, it is clear<br />
that this name was based on material <strong>of</strong> <strong>the</strong> common and widely distributed species generally known as<br />
Bacidia umbrina (Ach.) Bausch or Scoliciosporum umbrinum (Ach.) Arnold (basionym: Lecidea<br />
umbrina Ach, Lich. Univ. : 183 (1819)). In <strong>the</strong> 19th century 'syno<strong>the</strong>a' was commonly applied to Micarea<br />
denigrata. Despite Hedlund's comments, this misapplication has been continued by some authors, even in<br />
recent years (e.g. Ozenda & Clauzade, 1970: 402). To avoid <strong>the</strong> confusion that would arise if 'umbrina"<br />
were replaced by 'syno<strong>the</strong>a' , a formal proposal to reject <strong>the</strong> name Lecidea syno<strong>the</strong>a Ach. (and combinations<br />
from that name), under <strong>the</strong> provision <strong>of</strong> Art. 69.1, will be presented elsewhere.<br />
Thallus effuse and <strong>of</strong>ten widespreading, sometimes endoxylic but usually developing on <strong>the</strong><br />
surface <strong>of</strong> <strong>the</strong> substratum (especially in <strong>the</strong> vicinity <strong>of</strong> apo<strong>the</strong>cia) as convex to irregularly<br />
subglobose areolae. Areolae 60-200 ixm diam, greenish white to greenish grey, matt; in section<br />
without an amorphous covering layer, external hyphae hyaline but surrounding gel matrix <strong>of</strong>ten<br />
with dilute olivaceous, K-l- violet pigment. Thallus sometimes scurfy and dark grey-brown to<br />
blackish, owing to invasion by dematiaceous fungi and non-lichenized algae. Phycobiont<br />
micareoid, cells c. 4—7 /u,m diam.<br />
Apo<strong>the</strong>cia usually present and numerous (see 'observations' below), scattered to confluent,<br />
broadly convex to subglobose, sometimes tuberculate, 0- 15-0-5 mm diam, or to 0-6 mm when<br />
tuberculate; immarginate, or sometimes young apo<strong>the</strong>cia with an indistinct, shallow marginal<br />
rim, paler than <strong>the</strong> disc; disc pallid to brown or piebald (shade forms), more usually dark grey or<br />
black, matt. Hymenium (25-)30-40 /o-m tall, dilute olivaceous or dull brownish, K-l- violet;<br />
pigment <strong>of</strong>ten concentrated in upper part (with lower part ± hyaline), and confined to <strong>the</strong><br />
gel-matrix. Asci clavate, 28-36x9-12 jxm. Spores oblong-ellisoid, oblong-ovoid, fusiform or<br />
bacilliform, <strong>of</strong>ten slightly curved, (0-)l -septate, upper cell usually slightly shorter and broader<br />
than <strong>the</strong> lower, (7-)9-16(-18)x2-3-3(-3-5) /u,m. Paraphyses numerous, branched and sometimes<br />
anastomosing, l-l-5(-l-7) /i,m wide; apices scarcely wider, and never with closely<br />
adhering pigment. Hypo<strong>the</strong>cium 60-110 /xm tall, hyaline or very dilute yellowish straw; hyphae<br />
c. 1-1-5 /am wide, interwoven or some vertically orientated in upper part, intermixed with<br />
short-celled, ascogenous hyphae hyphae c. 2-4 /xm wide. Excipulum indistinct, but usually<br />
evident in sections <strong>of</strong> young, shallow-convex apo<strong>the</strong>cia, hyaline; hyphae radiating, branched<br />
and anastomosing, hyaline, c. 1-1-5 /xm wide.<br />
Pycnidia usually present and numerous, immersed in <strong>the</strong> thallus or substratum (endoxylic
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,
130 BRIAN JOHN COPPINS<br />
predominantly lignicolous species with small, black apo<strong>the</strong>cia, but all such species (except M.<br />
misella as discussed above) appear very different when examined microscopically.<br />
Habitat and distribution: M. denigrata is commonly found on <strong>the</strong> lignum <strong>of</strong> fallen trunks and<br />
old stumps <strong>of</strong> broad-leaved and coniferous trees, especially in ra<strong>the</strong>r open situations at <strong>the</strong> edges<br />
<strong>of</strong> woodlands, woodland glades, and hedgerows, etc. The communities in which it occurs are<br />
difficult to define, but to give some indication <strong>the</strong> following list <strong>of</strong> associated species has been<br />
made from collections from fallen decorticate trunks <strong>of</strong> Pinus in <strong>the</strong> native pine-woods <strong>of</strong><br />
Scotland: Buellia griseovirens, Cladonia coniocraea, C. macilenta, Hypogymnia physodes,<br />
Lecanora expallens, L. symmicta agg., Lecidea aeruginosa, L. icmalea, Micarea peliocarpa,<br />
Mycoblastus sterilis, Parmelia saxatilis, Platismatia glauca, Pseudevernia furfuracea, Xylographa<br />
abietina, and X. vitiligo.<br />
M. denigrata is a very successful primary coloniser <strong>of</strong> untreated timber-work. It also occurs on<br />
<strong>the</strong> same substrate whose preservative has lost its effectiveness, or whose painted surface has<br />
flaked <strong>of</strong>f. On such substrata M. denigrata has been found in a wide range <strong>of</strong> situations, e.g.<br />
gates, fence posts and rails, garden furniture, picnic tables and seats, window frames <strong>of</strong><br />
greenhouses, wooden ro<strong>of</strong> tiles (shingles), telegraph poles, and <strong>the</strong> woodwork <strong>of</strong> old carts and<br />
farm machinery. When on such worked wood it <strong>of</strong>ten forms almost pure stands, and is <strong>of</strong>ten<br />
present as a form with few apo<strong>the</strong>cia but numerous pycnidia containing mesoconidia; <strong>the</strong><br />
conidia <strong>of</strong>ten extruding as white blobs easily visible to <strong>the</strong> unaided eye or through a x 10 lens.<br />
Lichens associated with M. denigrata on worked wood in <strong>the</strong> <strong>British</strong> Isles include Buellia<br />
punctata, Cyphelium inquinans, Hypogymnia physodes, H. tubulosa, Lecanora conizaeoides, L.<br />
Map 7 Micarea denigrata # 1950 onwards O Before 1950
LICHEN GENUS MICAREA IN EUROPE 131<br />
dispersa agg., L. piniperda, L. pulicaris, L. saligna, L. symmicta agg., L. varia, Lecidea<br />
aeruginosa, L. icmalea, Mycoblastus sterilis, Parmelia sulcata, Scoliciosporum chlorococcum, S.<br />
umbrinum, Strangospora moriformis, and Thelocarpon laureri.<br />
Fur<strong>the</strong>r evidence <strong>of</strong> <strong>the</strong> pioneering abilities <strong>of</strong> M. denigrata comes from <strong>the</strong> find (Coppins<br />
1888) <strong>of</strong> it on hardboard lying in a dune-slack, and <strong>the</strong> observations by Poelt (1977; as M.<br />
hemipoliella) on its colonisation <strong>of</strong> dead leaves <strong>of</strong> Cladium mariscus in a fen in Bavaria. O<strong>the</strong>r<br />
reports on s<strong>of</strong>t vegetable matter are wanting, although I have found it on decaying mosses (with<br />
Cladonia chlorophaea agg.) on <strong>the</strong> slope <strong>of</strong> a stable sand dune on Holy Island in Northumberland<br />
{Coppins 4456).<br />
M. denigrata is occasionally found on <strong>the</strong> ra<strong>the</strong>r dry and loose bark <strong>of</strong> old trees (e.g. Acer<br />
pseudoplatanus, Alnus, Betula, Castanea, Sambucus, tjlmus, and PmM-s), usually towards <strong>the</strong>ir<br />
bases; however occurrences on ± smooth bark are very rare (e.g. type material <strong>of</strong> Lecidea<br />
hemipoliella, on Alnus in sou<strong>the</strong>rn Finland). To my knowledge M. denigrata never occurs on <strong>the</strong><br />
healthy, living twigs <strong>of</strong> trees, or shrubs (see 'observations' above). There are two <strong>British</strong><br />
collections on shaded sandstone, one from Fife in a conifer plantation, <strong>the</strong> o<strong>the</strong>r from Yorkshire<br />
in a ditch embankment; associated species included Lecidea granulosa agg. , L. icmalea, Micarea<br />
peliocarpa, and Parmeliopsis ambigua. In addition, it has been found growing with Psilolechia<br />
lucida on <strong>the</strong> east-facing vertical side <strong>of</strong> a tomb in a Suffolk churchyard.<br />
M. denigrata is widely distributed in mainland Britain, but has not yet been reported from<br />
Ireland. In mainland Europe it is again widespread although I have not seen material from north<br />
<strong>of</strong> about 67°N; its range extends into eastern Europe and <strong>the</strong> Balkans and continues to <strong>the</strong><br />
Caucasus. I have examined North American material from Newfoundland, Washington (state),<br />
and Colorado, thus suggesting a widespread distribution in that subcontinent.<br />
Exsiccata: Anzi Lich. Ital. 256 (BM). Arnold Lich. Mon. 46 (BM ex K). Britz. Lich. Exs. : 464<br />
p.p. (H).<br />
Fries Lich. Suec. 98 (E, UPS). Harm. Lich. Loth. 838 (ANGUC, DUKE). Hepp Flecht. Eur. 14 (E).<br />
Johnson Lich. Herb. 373 (BM). Krypt. Exs. Vindob. 3153 (BM, BM ex K), 3651 (BM, BM ex K, M), 4858<br />
(GZU). Kutak Lich. Bohem. 205 (O), 516 (O), 517 (O). Malbr. Lich. Norm. 387 (M). Malme Lich. Suec.<br />
145 (M, S). Migula Crypt. Germ. 132 (BM, E, MANCH). Mougeot & "HcsiXcx Stirpes Crypt. 1329p. min. p.<br />
(BM). Norrlin &. Nyl. Herb. Lich. Fenn. Ill (BM, H), 745 (BM, H). Poelt Lich. Alp. 22 (BM, M, WIS).<br />
Rabenh. Lich. Eur. 626 (BM, BM ex K, E). Samp. Lich. Port. 132 (LD). Vezda Lich. Sel. 1430 (BM).<br />
Weber Lich. Exs. 73 (E, DUKE, M, NMW, WIS). Zwackh Lich. Exs. 394 (BM ex K, H).<br />
12. Micarea elachista (Korber) Coppins & R. Sant. , comb. nov.<br />
(Figs IC, 14A)<br />
Biatora elachista Korber, Parerga lich.: 159 (1860). - Catillaria elachista (Korber) Vainio in Acta Soc.<br />
Fauna Fl. fenn. 57 (2): 455 (1934). Type: Germany, Baden-Wiirttemberg, Heidelberg, on old trunk <strong>of</strong><br />
Castanea sativa, W. E. von Ahles (L 910, 138-100-lectotype!; L 910, 138-32-isolectotype!) See note<br />
below.<br />
Lecidea anomala *L. glomerella Nyl., Lich. Scand.: 203 (1861). - Biatorina glomerella (Nyl.) Arnold in<br />
Flora, Jena 53: 474 (1870). - Catillaria glomerella (Nyl.) Th. Fr., Lich. Scand. 2: 578 (1874). - Micarea<br />
glomerella (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 75, 85 (1892). Type:<br />
Finland, Ostrobottnia australis, Botom, 1859, A. J. Malmgren (H-NYL 19123 - lectotype!).<br />
Lecidea poliococca Nyl., Lich. Scand. : 203 (1861). - Catillaria denigrata f. poliococca Vainio in Acta Soc.<br />
Fauna Fl. fenn. 57 (2): 460 (1934). - Catillaria syno<strong>the</strong>a var. poliococca (Nyl.) Erichsen in Annls mycol.<br />
41: 205 (1943). Type: Sweden, Uppland, 'Upsalia', in silva 'Parken', ad pinos decorticatos' , 1852, W.<br />
Nylander (H-NYL 19144- holotype!).<br />
Lecidea sororians Nyl. in Flora, Jena 58: 445 (1875). - Bacidia sororians (Nyl.) H. Olivier in Bull. Geogr.<br />
hot. 21: 168 (1911). Type: Finland, Tavastia australis, Korpilahti, near Raianlahti, on rock with<br />
Stigonema sp., 1873, E. A. Lang [Vainio] (H-NYL 17234 -holotype!).<br />
Lecidea glomerella f. simplicata Nyl. in Vainio in Medd. Soc. Fauna Fl. fenn. 10: 28 (1883). Type: Finland,<br />
Tavastia australis, Evo, 'supra truncum pineum', 1874, J. P. Norrlin, Norrlin & Nyl. Herb. Lich. Fenn.<br />
314 (H- lectotype!; isolectotypes: BM!,H!M!).<br />
Lecidea glomerella var. poliococcoides Vainio in Medd. Soc. Fauna Fl. fenn. 10: 29 (1883). Type: Finland,<br />
Karelia borealis, Lieksa, Vieki, on burnt lignum, 1875, E. A. Vainio (TUR-VAINIO 22326 - holov<br />
type!).
132 BRIAN JOHN COPPINS<br />
Catillaria elachista var. carbonicola Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 458 (1934). Type: Finland,<br />
Tavastia australis, Korpilahti, 'hiiltyneella kannolla', 1873, E. A. Lang [Vainio] (H-NYL 19143 -<br />
isotype!).<br />
Notes on <strong>the</strong> typification o/Biatora elachista Korber. In <strong>the</strong> protologue to B. elachista, Korber (loc. cit.)<br />
gives <strong>the</strong> following habitat and locality information: 'On alten Striinken der Castanea vesca bei Heidelberg<br />
von Hrn. v. Zwackh und Ahles aufgefunden.' Material borrowed from <strong>the</strong> Korber Herbarium (L)<br />
consisted <strong>of</strong> four specimens: i. bei Heidelberg, Zwackh, annotated: 'Biatora elachista Kbr. nov. sp.' (L<br />
910, 138-101). ii. Heidelberg, Ahles (L 910, 138-100). iii. identical label as ii. (L 910, 138-32). iv.<br />
Forstrevier Goleow bei Rybnik, iii 1872, B. Stein (L 910, 138-298). (i-iii) are on Castanea lignum and (iv) is<br />
on conifer lignum.<br />
The most obvious choice for lectotype is (i) but it is not a Micarea and does not conform to <strong>the</strong> usual<br />
interpretation <strong>of</strong> <strong>the</strong> name (e.g. Vainio, 1934: 45; Ozenda & Clauzade, 1970: 401). I do not know <strong>the</strong><br />
identity <strong>of</strong> this species, but it may be close to Lecidea hypopta Ach. ; a brief description follows:<br />
Thallus probably mostly endoxylic but obscured by a pale farinose crust <strong>of</strong> a non-lichenised alga.<br />
Apo<strong>the</strong>cia numerous, reddish brown to brown-black, epruinose 0- 15-0-3 mm diam. Young<br />
apo<strong>the</strong>cia with a pale margin which is soon excluded as <strong>the</strong> disc expands and becomes convex.<br />
Epi<strong>the</strong>cium reddish brown, turning olivaceous brown in K. Hymenium 30-35 /u,m tall. Paraphyses<br />
simple or forked in <strong>the</strong>ir upper part, c. 1-1-5 ^tm wide; upper 5-15 ^tm <strong>of</strong>ten with pigmented walls<br />
and up to 2-5 /x,m wide. Asci clavate 28-35x10-11 /xm, 8-spored. Spores simple or l-(rarely 2-)<br />
septate oblong-ellipsoid, straight or slightly curved, contents <strong>of</strong>ten becoming brown, 8-13x3-4 /u,m.<br />
Excipulum hyaline within, reddish brown at edge, <strong>of</strong>ten with a penetrating algal layer c. 25 )u,m wide.<br />
Algal cells c. 7-13 ^tm diam. Pycnidia not found.<br />
Specimens (ii) and (iii) are presumably those that Korber attributed to Ahles in <strong>the</strong> protologue and are<br />
<strong>the</strong>refore syntypes. Both belong to <strong>the</strong> species generally known as Catillaria elachista (Korber) Vainio, and<br />
it is with <strong>the</strong>se that <strong>the</strong> name is lectotypified here. Specimen (iv) is not a syntype and does not belong<br />
to ei<strong>the</strong>r <strong>of</strong> <strong>the</strong> above taxa. It has small, brown thinly white-pruinose apo<strong>the</strong>cia, a dark reddish<br />
brown hypo<strong>the</strong>cium and small, ellipsoid spores, 7-8x2-5-3 /am, and is referable to Lecidea apochroeella<br />
Nyl.<br />
Thallus effuse, superficial, consisting <strong>of</strong> dispersed to continuous, convex to subglobose<br />
areolae; areolae greenish white or whitish grey, sometimes tinged grey-brown or olivaceous,<br />
occasionally dark brown (when on burnt lignum), matt, sometimes ± white-pruinose, c.<br />
0-08-0-16(-0-25) mm diam. Areole in section (Fig. IC) with a c. 10-12 /u,m tall, hyaline or<br />
greyish (pigment in gel-matrix, K+ violet), algal-free 'cortex', composed <strong>of</strong> interwoven, hyaline<br />
hyphae (c. 1-5 ^tm wide) that separate in K; outer surface <strong>of</strong> cortex sometimes bound by a<br />
hyaline, amorphous, densely gelatinised layer ('epicortex'), c. 3-5 /xm tall. This organised<br />
structure with a 'cortex' and 'epicortex' is <strong>of</strong>ten disrupted by <strong>the</strong> invading torulose hyphae <strong>of</strong> a<br />
dematiaceous hyphomycete. Phycobiont micareoid, cells 4-7 jxm diam.<br />
Apo<strong>the</strong>cia usually numerous, immarginate, convex to ± globose, <strong>of</strong>ten becoming tuberculate,<br />
dark brown to brown-black, matt, (0-08-)0- 12-0-3 mm diam, or to 0-8 mm diam when<br />
tuberculate. Hymenium 30-40 />tm tall. Upper part (epi<strong>the</strong>cium) usually well defined, up to<br />
12 jxm tall, dark fuscous-brown; pigment concentrated into dense amorphous clumps, but<br />
dissolving and fading into solution (without changing colour) in K, HNO3— and not dissolving;<br />
epi<strong>the</strong>cium <strong>of</strong> young apo<strong>the</strong>cia sometimes Kf+ violet due to <strong>the</strong> additional presence <strong>of</strong> <strong>the</strong> dull<br />
olivaceous, K-l- violet pigment which occurs in greater concentrations in <strong>the</strong> pycnidia. Remain-<br />
ing (lower) part <strong>of</strong> hymenium, hyaline with dilute yellowish brown vertical streaks. Asci clavate,<br />
23-35x10-12 jxm. Spores fusiform, oblong-fusiform or ovoid-oblong, <strong>of</strong>ten slightly curved,<br />
mostly 0-1-septate and slightly constricted at <strong>the</strong> septum, (9-)ll-15(-19)x2-3'5 jxm; occasionally<br />
becoming 2- or 3-septate and up to 20-24 /xm long; old spores sometimes with brownish<br />
contents. Paraphyses numerous, hyaline throughout, 0-8-1 /u,m wide in mid-hymenium but<br />
gradually widening above to l-7(-2) /xm; sparingly branched and sometimes anastomosing, but<br />
becoming richly branched above where <strong>the</strong>ir entangled apices, toge<strong>the</strong>r with <strong>the</strong> brown<br />
pigment, form a ± well delimited epi<strong>the</strong>cium. Hypo<strong>the</strong>cium 60-150 /xm tall, pale, tinged dilute<br />
yellowish brown, K— , HNO3-; hyphae hyaline, 1-1-5 /xm wide, interwoven, becoming ±<br />
vertically orientated towards <strong>the</strong> hymenium; ascogenous hyphae with swollen cells, mostly 2-
LICHEN GENUS MICAREA IN EUROPE 133<br />
4 fim wide. Excipulum poorly developed and much reflexed, sometimes evident as a narrow,<br />
pale fuscous-brown zone; hyphae hyaline, radiating, branched and anastomosing, 1-1-5 /xm<br />
wide.<br />
Pycnidia usually present and numerous, developing from within areolae but soon becoming<br />
emergent and sometimes ± sessile; <strong>of</strong> two types: (a) c. 100-200 /xm diam, grey-brown but<br />
usually paler or whitish around <strong>the</strong> ostiole, surface smooth and ± glossy; ostioles distinct and c.<br />
20 jxm diam, sometimes gaping and to 50 jxva diam; wall not continuing below <strong>the</strong> base, laterally<br />
c. 23-40 /xm wide, dilute olivaceous or brownish, K-l- violet, and formed <strong>of</strong> interwoven hyaline<br />
hyphae (c. 1-5 /xm wide) that ± separate in K; conidiogenous cells cylindrical, occasionally<br />
percurrently proliferating, c. 4—7x1-2 /xm; conidia (mesoconidia) ± cylindrical, sometimes<br />
slightly wider at proximal end, <strong>of</strong>ten faintly biguttulate, 3-5-4-5xl-3-l-7(-2) /xm. (b) similar in<br />
appearance and structure to above, but smaller and 60-100 /xm diam; conidiogenous cells<br />
cylindrical, c. 3-6x1 /xm; conidia (microconidia) narrowly cylindrical, (4-)4-5-6(-6-5)x<br />
0-7-1 /xm.<br />
Chemistry: Thallus K— , C—<br />
by t. I.e.<br />
, KC— , PD— ; sections <strong>of</strong> apo<strong>the</strong>cia C— ; no substances detected<br />
Observations: The combination <strong>of</strong> ± globose areolae, ± globose, brown apo<strong>the</strong>cia, dense<br />
brown (dissolving in K) epi<strong>the</strong>cial pigment, C- hymenium, fusiform, 1-3-septate spores and<br />
distinctive, ± glossy pycnidia usuall-y make this species easy to identify. Some confusion has<br />
been made with shade forms <strong>of</strong> M. denigrata, but <strong>the</strong>se can be distinguished by <strong>the</strong>ir ra<strong>the</strong>r<br />
adnate and <strong>of</strong>ten larger apo<strong>the</strong>cia, C-f- orange-red (gyrophoric acid) hymenium and thallus,<br />
never glossy, thin-walled pycnidia, and, when present, curved macroconidia. M. elachista is<br />
closely related to M. rhabdogena (q-v.), which differs in having an endoxylic thallus, smaller,<br />
mostly simple, spores, and black pycnidia.<br />
The areolae <strong>of</strong> M. elachista sometimes have a white-pruinose appearance: <strong>the</strong> 'pruina'<br />
resulting from <strong>the</strong> partial disintegration <strong>of</strong> <strong>the</strong> thin 'epicortex'.<br />
Habitat and distribution: M. elachista is found, <strong>of</strong>ten in <strong>the</strong> company <strong>of</strong> Parmeliopsis spp. , on<br />
<strong>the</strong> lignum (rarely bark) <strong>of</strong> partially or wholly decorticate trunks or large stumps <strong>of</strong> old trees,<br />
especially Castanea, Pinus, and Quercus. Most collections from France and Germany were<br />
made from Castanea, whereas those made from Scandinavia were mostly from conifers. It<br />
occasionally occurs on burnt or charred stumps and my collection (Coppins 6017) from Sweden,<br />
on Pinus, was accompanied by Chaeno<strong>the</strong>ca ferruginea, Hypocenomyce friesii, Lecidea granulosa<br />
agg. , Micarea melaena, and Parmeliopsis spp. It seems to be rare on worked timber, but was<br />
collected on old fence-posts in Bavaria by Arnold.<br />
I have seen only one saxicolous ga<strong>the</strong>ring <strong>of</strong> M. elachista, i.e. <strong>the</strong> holotype <strong>of</strong> Lecidea<br />
sororians from sou<strong>the</strong>rn Finland, with which occurred colonies <strong>of</strong> a Stigonema sp. , a few lobes <strong>of</strong><br />
a brown Parmelia (? P. verruculifera) and, according to Nylander, Biatorella torvula (not<br />
present in <strong>the</strong> existing material). This specimen is very small and in poor condition, and <strong>the</strong> few<br />
spores seen appeared to be abnormally developed, 1-3-septate, 15-24x2-2-5 /xm; o<strong>the</strong>r<br />
features <strong>of</strong> <strong>the</strong> thallus and apo<strong>the</strong>cia agree with M. elachista (pycnidia were not found) and I<br />
think it is most unlikely to represent a distinct taxon.<br />
M. elachista appears to avoid <strong>the</strong> more strongly oceanic areas <strong>of</strong> Europe, and is known from<br />
mid-Sweden, sou<strong>the</strong>rn Finland, France (Haute Vienne, <strong>the</strong> Massif Central and <strong>the</strong> Pyrenees),<br />
sou<strong>the</strong>rn Germany (Hessen, Baden-Wiirttemberg and Bavaria), and <strong>the</strong> Austrian Tirol. Its<br />
presence in Britain is a possibility and it should be sought for, especially in <strong>the</strong> central Highlands<br />
and east Scotland, and <strong>the</strong> Welsh border counties.<br />
Exsiccata: Arnold Lick. Exs. 1471 (BM ex K, M). Arnold Lick. Mon. 246 (BM ex K, M). Malme Lick.<br />
Suec. 21 (M, S). Norrlin & Nyl. Herb. Lick. Fenn. 314 (BM, H, M); 724 (BM). Vezda Lick. Sel. 1134<br />
(BM). Zwackh Lich. Exs. 122 (H-NYL 18828, M).
134 BRIAN JOHN COPPINS<br />
13. Micarea eximia Hedl.<br />
(Figsl4B,40C)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 80, 84, 95 (1892). - Catillaria malmeana Zahlbr.,<br />
Cat. lich. univ. 4: 56 (1926); nom. nov.; non Catillaria eximia Malme. Type: Sweden, Dalarna, Alvdal,<br />
Hallstugen, vi 1891, J. T. Hedlund (S - lectotype! [t.l.c: no substances]; S - isolectotype!; material<br />
distributed as Malme Lich. Suec. Exs. 26 is possibly part <strong>of</strong> this collection).<br />
Thallus effuse, endoxylic, <strong>of</strong> minute clusters (c. 15-40 yun diam) <strong>of</strong> <strong>the</strong> phycobiont amongst<br />
<strong>the</strong> wood fibres, with intertwining hyphae that <strong>of</strong>ten have green walls, K-, HNO3+ red.<br />
Phycobiont micareoid, cells 4-7 /x,m diam.<br />
Apo<strong>the</strong>cia numerous, immarginate, ± convex-globose from <strong>the</strong> start, <strong>of</strong>ten becoming<br />
tuberculate, black, matt or slightly glossy, 0- 1-0-2 mm diam, or to 0-4 mm diam when<br />
tuberculate. Hymenium 30-45 /am tall; upper part bright to dark aeruginose, K— or dulling,<br />
HNO3+ purple-red; remaining (lower) part ± hyaline with aeruginose vertical streaks. Asci<br />
clavate, c. 30-40x11-12 ^.m. Spores oblong-fusiform, sometimes slightly curved, 9-14<br />
(-16) X 1 •8-2-5 /xm. Paraphyses ra<strong>the</strong>r scanty, branched and anastomosing, thin and 0-7-0-8 /xm<br />
wide, but upper c. 5-17 (xm with thickened, pigmented walls and <strong>the</strong>n 2-3 /xm wide; usually a few<br />
paraphyses have pigmented walls throughout <strong>the</strong>ir length and are about 2 ^im wide in <strong>the</strong> middle<br />
hymenium. Hypo<strong>the</strong>cium 45-120 //,m tall, dilute reddish- or purplish brown, sometimes with<br />
darker blotches, K+ dull greenish, HNO3+ purple-red, pigment confined to gel-matrix; hyphae<br />
interwoven, hyaline, c. 1-1-5 /xm wide; ascogenous hyphae with swollen cells up to 4 ^im diam.<br />
Excipulum indistinct, sometimes evident as a dark green or brownish (K-l- olivaceous) reflexed<br />
zone with branched, radiating, pigmented hyphae c. 1-5-2 /xm wide.<br />
Pycnidia numerous but inconspicuous, usually immersed between surface wood fibres, but<br />
sometimes ± emergent, black, c. 35-80 /xm diam; walls dark green, K-, HNO3-I- purple-red,<br />
composed <strong>of</strong> tightly bound, pigmented hyphae, c. 2 /xm wide. Conidia (mesoconidia) cyhndri-<br />
cal, sometimes biguttulate, 3-9-5-5X 1-1-4 /xm.<br />
Chemistry: Sections <strong>of</strong> apo<strong>the</strong>cia and thallus C— ; no substances detected by t.l.c.<br />
Observations: Micarea eximia is characterized by <strong>the</strong> combination <strong>of</strong> an endoxylic thallus,<br />
small, black, ± globose or tuberculate apo<strong>the</strong>cia, bright green upper hymenium, reddish or<br />
purplish brown (K-l- green) hypo<strong>the</strong>cium, simple or 1-septate, oblong-fusiform spores, and an<br />
absence <strong>of</strong> stalked pycnidia. It is most likely to be confused with M. contexta and M. olivacea.<br />
The former has broader, ovoid spores and a darker hypo<strong>the</strong>cium; and <strong>the</strong> latter has a less<br />
brightly coloured hymenium, a darker olivaceous hypo<strong>the</strong>cium, more numerous and broader<br />
paraphyses, shorter and slightly broader spores with rounded apices, and shorter mesoconidia.<br />
M. nigella differs from M. eximia in having a purplish brown upper hymenium, a darker<br />
hypo<strong>the</strong>cium (hyphal walls pigmented), ellipsoid to oblong-ovoid, simple spores, and stalked<br />
pycnidia. The spores <strong>of</strong> M. eximia are similar to those <strong>of</strong> M. denigrata, but that species has an<br />
olivaceous, K-l- violet pigment in its upper hymenium and pycnidial walls, and a ± hyaline<br />
hypo<strong>the</strong>cium.<br />
Habitat and distribution: M. eximia is a rare or overlooked species <strong>of</strong> conifer lignum, known<br />
only from middle Sweden and nor<strong>the</strong>rn Finland. Associated species on <strong>the</strong> specimens examined<br />
include Bacidia retigena, Calicium glaucellum, Cetraria pinastri, Cladonia spp., Lecideapullata,<br />
Micarea misella, Parmeliopsis aleurites, P. ambigua, P. hyperopia, and Xylographa vitiligo.<br />
14. Micarea globulosella (Nyl.) Coppins, comb. nov.<br />
(Figsl5,43A-B;Map24)<br />
Lecidea globulosella Nyl., Lich. Jap.: 69 (1890). - Bacidia globulosella (Nyl.) Zahlbr., Cat. lich. univ. 4:<br />
202 (1926). Type: Japan, Yokohama, on bark, 1879, E. /4/m^Mwf(S -lectotype! ; isolectotypes: H-NYL<br />
17412! and 17413!, S!).<br />
Thallus effuse, <strong>of</strong> scattered or, more usually, ± contiguous areolae. Areolae convex, whitish<br />
or grey, not gelatinous when wet, 40-150 /xm diam; in section without an amorphous covering
LICHEN GENUS MICAREA IN EUROPE 135<br />
layer, outer hyphae sometimes olivaceous (or brownish) and K+ violet. Phycobiont micareo'id,<br />
cells 4-7 /Ltm diam.<br />
Apo<strong>the</strong>cia numerous, immarginate, convex and adnate to subglobose, sometimes becoming<br />
tuberculate, greyish or brownish black, sometimes pale grey or pallid (shade forms), 0- 1-0-3 mm<br />
diam. Hymenium 35-40 /xm tall, dilute olivaceous or dilute olive-brown, K+ violet. Asci<br />
clavate, 30-35x9-12 jxm. Spores fusiform-acicular or ± rod-shaped, slightly curved or ±<br />
straight, 0-3(-6)-septate, 13-26 x(l'5-)2-2-5(-3) /xm. Paraphyses numerous, branched and<br />
sometimes anastomosing, c. 1 fxm wide, sometimes widening to 1-5 fxm towards <strong>the</strong>ir apices;<br />
apical walls hyaline. Hypo<strong>the</strong>cium 50-70 /xm tall, hyaline. Excipulum indistinct, evident in some<br />
sections as a narrow, reflexed, lateral border to <strong>the</strong> hymenium, hyaline or dull olivaceous (<strong>the</strong>n<br />
K-l- violet); hyphae radiating, branched and anastomosing, c. 1 jxm wide.<br />
Pycnidia frequent but inconspicuous, immersed or emergent, whitish to dark grey; walls<br />
olivaceous or dull brownish, K-l- violet (especially around <strong>the</strong> ostioles), sometimes ± hyaline.<br />
Pycnidia <strong>of</strong> two types: (a) immersed within large areolae, sometimes emergent, 60-100 />im<br />
diam; conidia (mesoconidia) ± cylindrical, 3-6-5-3X 1-1-4 /xm; (b) similar but smaller, c. 30-40<br />
fxm diam; conidia (microconidia) narrowly cylindrical, 3-8-5x0-8-1 /am.<br />
Chemistry: Thallus C+ red, K— , PD—<br />
to oHvaceous pigment); t.l.c: gyrophoric acid.<br />
; apo<strong>the</strong>cia sections C— or C-l- red (also C-l- violet due<br />
Observations: M. globulosella is closely related to M. denigrata and M. nitschkeana but can be<br />
distinguished by its longer, almost acicular, or rod-shaped spores. My early suspicions that M.<br />
globulosella could be a long-spored variant <strong>of</strong> M. nitschkeana were removed by <strong>the</strong> examination<br />
<strong>of</strong> microconidia, those <strong>of</strong> M. nitschkeana being significantly longer (mostly 5-5-7 /am). Long,<br />
curved macroconidia, as found in M. denigrata and M. nitschkeana, have not yet been found in<br />
M. globulosella, but <strong>the</strong>y should be sought for in additional material. See under M. syno<strong>the</strong>oides<br />
for differences from that species and Bacidia beckhausii. The name Micarea bacidiella (a<br />
synonym <strong>of</strong> B. beckhausii) was mistakenly applied to M. globulosella by Vezda & Wirth (1976)<br />
and Poeh&Vezda (1977).<br />
Habitat and distribution: M. globulosella is a rare but widespread species, so far known to me<br />
from Wales, Sweden, Finland, south-east France, Bavaria (Allgau), Czechoslovakia (High<br />
Tatra) , Canada (Quebec) , and Japan (Honshu) . It seems to prefer conifer bark , but at <strong>the</strong> Welsh<br />
locality it occurred on <strong>the</strong> top <strong>of</strong> an old gate in a wooded valley, and was accompanied by<br />
Lecanora piniperda, Lecidea aeruginosa, and L. icmalea. The specimen from Allgauer Alpen is<br />
associated with Graphis scripta, Menegazzia terebrata, Stenocybe major, and Frullania sp. on<br />
Abies; that from Sweden is on Picea with Cetraria pinastri and Ptilidium pulcherrimum; and<br />
those from Czechoslovakia are on Picea, with Cetraria chlorophylla, Hypogymnia physodes,<br />
and Ptilidium pulcherrimum. At <strong>the</strong> French locality it was growing on an old basidiome <strong>of</strong> <strong>the</strong><br />
polypore Daedaleopsis confragosa on Pinus hapelensis.<br />
It seems to require humid conditions in old forest situations, but has a more widespread<br />
distribution than M. syno<strong>the</strong>oides which is strongly oceanic. M. globulosella may have a greater<br />
tolerance to cold winter temperatures, such as occur in <strong>the</strong> central European montane regions.<br />
Exsiccata: Rasanen Lichenoth. Fenn. 426 (hb Vezda).<br />
15. Micarea hedlundii Coppins, sp. nov.<br />
(Figs 14C, 44A)<br />
Thallus effusus, olivaceo-viridis, subtiliter granulosus ad 0-4 mm crassus, constatus ex goniocystis;<br />
goniocystae c. 20-40 /u.m diam, omnes cum pigmento flavo-brunneolo, K+ purpureo-violaceo et oleoso.<br />
Algae cellulis 4-7 /u,m diam. Apo<strong>the</strong>cia vulgo pauca vel etiam nulla, immarginata, convexo-hemisphaerica<br />
mox tuberculata, pallida vel griseo-fusca demum obscure fusca, 0-15-0-5 /xm diam. Hymenium c. 35 /xm<br />
altum, ± hyalinum vel p.p. dilute olivaceo-brunneolum, K-l- violaceum. Ascosporae ellipsoideae,<br />
ovoideae vel oblongae, simplices interdum 1- septatae, 6-5-10(-12)x 2-3-4 /xm. Paraphyses aliquantum<br />
paucae, ramosae, c. 0-7-l(-l-5) /xm latae, apicibus vix incrassatis, incoloratis, Hypo<strong>the</strong>cium pallidum.
136 BRIAN JOHN COPPINS<br />
Excipulum paulum evolutum. Pycnidia numerosa, conspicua, stipitata, 0- 1-1-0 mm alta, 0-07-0-14 mm<br />
diam, cum stipibus simplicibus vel ramosis, toto griseo-brunneo vel roseo-brunneo cum tomento exili<br />
albido. Conidia oblongo-ellipsoidea vel oblongo-ovoidea, (4-)4-5-5-5(-6)xl-3-l-7 ^tm.<br />
Typus: Norvegia, Opplandia, par. Ringebu, ad oriento-boreo-orientum ex Ringebu, ad S0raa, inter<br />
Nyhamnsbekken et Ulveslabekken, alt. 400 m, ad truncum decorticatum, 25 viii 1979, leg. L. Tibell 8657<br />
(UPS-holotypus).<br />
Thallus epixylic or over bryophytes on lignum, effuse, dull olive-green, consisting <strong>of</strong> fine<br />
granules (goniocysts) forming a loose crust to c. 0-4 mm thick. Goniocysts c. 20-40 /xm diam, <strong>the</strong><br />
centre <strong>of</strong> each with a yellow-brown or dull orange pigment which in K appears as purple-violet<br />
oily droplets. Phycobiont micareoid, cells 4-7 /um diam.<br />
Apo<strong>the</strong>cia few or sometimes absent, immarginate, convex and soon becoming tuberculate,<br />
pallid to grey-brown or dark brown, 0-15-0-5 mm diam. Hymenium c. 35 /xm tall, ± hyaline with<br />
vertical streaks <strong>of</strong> straw-brown or pale olive-brown, K+ violet, C+ violet, HNO3-I- reddish;<br />
lower hymenium sometimes with dull orange, K+ purple-violet (oily droplets) pigment. Asci<br />
clavate, c. 30x12 ixm. Spores ellipsoid, ovoid or oblong, simple, a few sometimes 1-septate,<br />
6-5-10(-12)x2-3-4 /am. Paraphyses ra<strong>the</strong>r scanty, branched and sometimes anastomosing,<br />
0-7-l(-l -5) /Ltm wide, apices not swollen or pigmented. Hypo<strong>the</strong>cium c. 70-100 /xm tall, hyaline,<br />
or with dull orange, K-f- purple-violet (oily droplets) pigment in upper part; hyphae interwoven,<br />
becoming vertically orientated towards <strong>the</strong> hymenium, c. 1-1-7 (xm wide, intermixed with<br />
swollen short-celled ascogenous hyphae. Excipulum indistinct and soon reflexed, sometimes<br />
discernible in young apo<strong>the</strong>cia as a narrow, hyaline, non-amyloid zone; hyphae radiating,<br />
branched and anastomosing, c. 0-8-1 /zm wide.<br />
Pycnidia numerous and conspicuous, stalked with one or to about five borne terminally on<br />
simple or branched pycnidiophores. Pycnidiophores (including pycnidia) grey-brown or pinkish<br />
brown, covered with a thin whitish tomentum, c. 01-1 mm tall and 0-07-0-14 mm diam; lower<br />
parts <strong>of</strong> pycnidophores <strong>of</strong>ten covered in goniocysts. Pycnidiophore and pycnidial wall tissues<br />
composed <strong>of</strong> hyaline hyphae bound by a dilute reddish brown matrix reacting K+ violet or<br />
violet-brown and HNO3-I- reddish; surface <strong>of</strong> pycnidiophores and pycnidia with protruding,<br />
slender, flexuose, hyaline tomental hyphae c. 0-7-1 /xm wide. Conidiogenous cells ampuUiform<br />
to ± cylindrical, 5-lOxc. 1-5 /am, <strong>of</strong>ten swollen at base to 2-5 /am wide. Conidia (mesoconidia)<br />
oblong-ellipsoid or oblong-ovoid, (4-)4-5-5-5(-6)xl-3-l-7 /am.<br />
Chemistry: Thallus C— , PD—<br />
; apo<strong>the</strong>cia sections C— (but with C+ violet pigment); no<br />
substances or traces <strong>of</strong> 'prasina-unknown B' (? contaminant) detected by t.l.c.<br />
Observations: Micarea hedlundii is readily identified by its finely granular, darkish green<br />
thallus with distinctly stalked, pinkish brown, tomentose pycnidia. Several o<strong>the</strong>r Micarea<br />
species have stalked pycnidia but in none <strong>of</strong> <strong>the</strong>m are <strong>the</strong>y tomentose. Ano<strong>the</strong>r unique feature<br />
oiM. hedlundii is <strong>the</strong> dull orange pigment, present in <strong>the</strong> goniocysts and sometimes in <strong>the</strong> lower<br />
hymenium and upper hypo<strong>the</strong>cium, which appears as purple-violet oily droplets in K. Unlike<br />
<strong>the</strong> olivaceous K-f violet pigment found in this species and o<strong>the</strong>rs (e.g. M. denigrata, M.<br />
nitschkeana, and M. prasina), it remains unchanged in C and 50% HNO3. M. hedlundii is<br />
unlikely to be confused with any o<strong>the</strong>r Micarea, but is probably closely related to M. prasina. M.<br />
anterior has reddish brown, stalked pycnidia but <strong>the</strong>y are glabrous and produce shorter conidia;<br />
in addition its apo<strong>the</strong>cia and pycnidia are completely devoid <strong>of</strong> K+ violet pigments.<br />
The chemistry <strong>of</strong> M. hedlundii is problematical. Of <strong>the</strong> three specimens tested by t.l.c, <strong>the</strong><br />
type from Norway contains no detectable substances, but <strong>the</strong> two specimens from Austria<br />
(GZU) and Germany (hb Poelt) appear to contain small amounts <strong>of</strong> 'prasina-unknown B'. This<br />
substance is known elsewhere only in <strong>the</strong> type race <strong>of</strong> M. prasina and it is possible that its<br />
detection in <strong>the</strong> aforesaid samples is due to contamination by that species. However, <strong>the</strong><br />
production <strong>of</strong> 'prasina-unknown B' by M. hedlundii is a possibility meriting fur<strong>the</strong>r study,<br />
especially as M. hedlundii and M. prasina appear to be closely related to one ano<strong>the</strong>r. New and<br />
carefully collected specimens <strong>of</strong>M hedlundii are required to resolve this problem.<br />
Habitat and distribution: M. hedlundii occurs on old stumps (? mainly <strong>of</strong> conifers) in<br />
woodlands. Few associated lichens are present on <strong>the</strong> specimens examined, mostly just a few
LICHEN GENUS MICAREA IN EUROPE 137<br />
scattered squamules <strong>of</strong> Cladonia spp. and fragments <strong>of</strong> Lepraria spp, although one <strong>of</strong> <strong>the</strong><br />
Norwegian collections includes a Chaeno<strong>the</strong>copsis. M. hedlundii seems to be a very rare species,<br />
but is known from scattered localities in Norway, Sweden, Germany (Bayern), Austria<br />
(Steiermark), and possibly Switzerland.<br />
Etymology: This new species is named in honour <strong>of</strong> Johan Teodor Hedlund (1861-1953) in<br />
recognition <strong>of</strong> his pioneering study <strong>of</strong> Micarea included in his doctorate <strong>the</strong>sis for <strong>the</strong> University<br />
<strong>of</strong> Uppsala (Hedlund, 1892), a work that has been a constant inspiration during my own studies.<br />
16. Micarea incrassata Hedl.<br />
(Figs4C, 16,41A-B;Map3)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 82, 94 (1892). Type: Austria, 'supra muscos in<br />
summo jungo Kraxentrag circa Brenner, Tirol, c. 2800 m. alt.', A. Minks (S - holotype!).<br />
Lecidea assimilata B. [var.] infuscata Th. Fr., Lich. Scand. 2: 522 (1874). - Lecidea assimilata f. infuscata<br />
(Th. Fr.) Vainio in Medd. Soc. Fauna Fl. fenn. 10: 85 (1883). Type: Norway, Sor-Trondelag, Oppdal<br />
hd., Dovre, Kongsvoll, Hogsnyta, 17 viii 1863, Th. M. Fries (UPS - lectotype!; UPS - isolectotype!<br />
[t.l.c: no substances]).<br />
Thallus growing on bryophytes, plant debris or sandy soil, composed <strong>of</strong> confluent, convexverrucose<br />
areolae that are intermixed with cephalodia. Areolae dull grey-white, or grey-brown<br />
to dark grey, matt, 0-08-0-3 mm diam; in section, usually without a hyaline amorphous covering<br />
layer (but sometimes seen in sections <strong>of</strong> young areolae), hyphae in outer c. 10 jxm <strong>of</strong>ten with<br />
light brown walls. A ± algae-free medulla <strong>of</strong>ten differentiated. Thallus hyphae c. T8-3 /am<br />
wide. Phycobiont micareoid, cells 4—7 /u,m diam. Cephalodia <strong>of</strong>ten present, irregularly globose<br />
and hidden amongst <strong>the</strong> areolae but sometimes visible externally as brown areolae-like<br />
structures, 0-2-0-6 mm diam; containing Nostoc, cells 3-5 /i,m diam. Less <strong>of</strong>ten present are<br />
irregular, ra<strong>the</strong>r loose clusters (? cephalodia) <strong>of</strong> Stigonema.<br />
Apo<strong>the</strong>cia numerous, immarginate, convex, ± adnate and <strong>of</strong>ten partly immersed by surrounding<br />
areolae, black, matt, 0-3-0-8(-l) mm diam, sometimes forming irregular tuberculate<br />
clusters up to T2 mm diam. Hymenium 45-50 ^im tall; upper c. 10 ixm (epi<strong>the</strong>cium) dark<br />
aeruginose or olivaceous, K— , HNO3-I- red; remaining (lower) part dilute greenish or hyahne.<br />
Asci clavate, 45-48x11-14 [xm. Spores ellipsoid, oblong-ellipsoid, oblong-ovoid or oblongfusiform,<br />
0-l(-2)-septate, (10-)12-17x4-4-8 /xm. Paraphyses numerous, simple below, sometimes<br />
forked above, sometimes anastomosing, (l-)l-5-2 ixm wide, sometimes widening above<br />
to 3 jxm; apical walls hyaline although surrounded by dense pigment in <strong>the</strong> surrounding gel<br />
matrix. Hypo<strong>the</strong>cium c. 150-400 /xm tall, sometimes 'rooting' to <strong>the</strong> base <strong>of</strong> <strong>the</strong> adjoining<br />
areolae, dark red-brown, without a purple tinge, K-, HNO3+ bright orange-brown; hyphae<br />
interwoven, or ± vertically arranged in upper part, c. T7-2-5(-3) ^tm wide, surrounded by<br />
densely pigmented matrix; ascogenous hyphae c. 2-5-5 /xm wide. Excipulum indistinct,<br />
sometimes evident as a reflexed, dull olivaceous or reddish brown zone; hyphae radiating,<br />
branched and anastomosing, c. 1-2 /xm wide.<br />
Pycnidia rare, immersed to sessile, black, 30-60 /am diam; walls dark green above, changing<br />
to reddish brown at base; conidia (? mwroconidia) cylindrical or cylindrical-fusiform, 6-9x1-<br />
T3/am.<br />
Chemistry: Thallus K-, C-, PD-; t.l.c: no substances.<br />
Observations: See M. assimilata.<br />
Habitat and distribution: M. incrassata occurs in much <strong>the</strong> same habitats as M. assimilata but<br />
appears to be more widely distributed. In Europe it has an arctic-alpine distribution ranging<br />
from Spitzbergen (c. 79°N) in <strong>the</strong> high arctic, southwards to <strong>the</strong> Kola peninsula, central Norway<br />
(Opland), and central Sweden (Harjedalen). Fur<strong>the</strong>r south it occurs in <strong>the</strong> Scottish highlands<br />
(Clova in Angus, and near Loch Merkland in East Su<strong>the</strong>rland), Denmark, and <strong>the</strong> Alps (Upper<br />
Bavaria, Austrian Tirol, and Switzerland). I have not seen it from Alaska or Canada although it<br />
probably occurs in those regions. From fur<strong>the</strong>r south in North America it has been collected at
138 BRIAN JOHN COPPINS<br />
altitudes <strong>of</strong> c. 3400-3900 m in <strong>the</strong> Rocky Mountain National Park in Colorado by Anderson<br />
(1964). M. incrassata has been collected on Kerguelen Island in <strong>the</strong> sou<strong>the</strong>rn Indian Ocean<br />
(48°30'S) and is so far <strong>the</strong> only arctic or arctic-alpine Micarea known to have a bipolar<br />
distribution.<br />
Exsiccata: Fellman Lich. Arct. 164 (BM ex K, H, H-NYL 16567), 166 (BM ex K, H).<br />
17. Micarea intrusa (Th. Fr.) Coppins & Killias, comb. nov.<br />
(Figs 17, 55; Map 22)<br />
Lecidea intrusa Th. Fr. in Bot. Notiser 1867: 152 (1867). - Catillaria intrusa (Th. Fr.) Th. Fr., Lich. Scand.<br />
2: 579 (1874). - Lecidiopsis intrusa (Th. Fr.) Zopf in Hedwigia 35: 338 (1896). - Conida intrusa (Th. Fr.)<br />
Sacc, Syll. Fung. 18: 187 (1906). Type: Finland, Tavastia australis, Mustiala, 1867, A. Kullhem (UPS-<br />
holotype!).<br />
Lecidea aphanoides Nyl. in Flora, Jena 51: 476 (1868). Type: Scotland, South Aberdeenshire, Braemar,<br />
'Craig Guie', viii 1868, J. M. Crombie (H-NYL 20237 - holotype!); supposed isotype material in BM<br />
appears to be all Scoliciosporum umbrinum (Ach.) Arnold.<br />
Lecidea melaphana Nyl. in Flora, Jena 52: 83 (1869). Type: Scotland, South Aberdeenshire, Braemar,<br />
'Craig Guie', viii 1868, J. M. Crombie (BM - isotype!).<br />
Lecidea contrusa Vainio in Medd. Soc. Fauna Fl. fenn. 10: 29 (1883); nom. illeg. (Art. 63).<br />
Thallus occasionally effuse but more <strong>of</strong>ten forming small patches (to c. 1 cm diam) amongst<br />
o<strong>the</strong>r lichens, irregularly granular-verrucose and <strong>of</strong>ten rimose, sometimes forming discrete<br />
areolae (c. 0- 1-0-3 mm diam) which may coalesce to form 'composite areolae' up to c. 1 mm<br />
diam and 0-5 mm tall. Thallus dark olive-grey or brownish grey; surface matt, and usually scurfy<br />
due to invasion <strong>of</strong> free-living algae. In section, sometimes with a hyaline amorphous covering<br />
layer up to 12 /xm thick but this is usually disrupted by invading algae ; walls <strong>of</strong> uppermost hyphae<br />
sometimes thickened with olive-green pigment, K-, HNO3-I- red; phycobiont layer c. 70^100<br />
mm thick, above a ± distinct hyaline medulla. Phycobiont not micareoid; cells globose, large,<br />
7-21 /xm diam, with thick hyaline walls 1-2 /xm thick, each cell deeply penetrated by a<br />
haustorium <strong>of</strong> <strong>the</strong> mycobiont (Fig. 55).<br />
Apo<strong>the</strong>cia numerous, immarginate, convex, adnate, black, or slightly glossy when young,<br />
0-14-0-4 mm diam. Hymenium 40-50 /xm tall; upper (c. 20 /x,m) part aeruginose-green, K-,<br />
HNOs-t- red; remaining (lower) part ± hyaline. Asci clavate, 35-45x14-15 /xm; cytoplasm <strong>of</strong><br />
asci and spores sometimes with an orange, K+ purple-red pigment. Spores elhpsoid or<br />
ovoid-ellipsoid, occasionally oblong-ellipsoid or oblong fusiform and <strong>the</strong>n sometimes slightly<br />
curved, simple or 1-septate and septum <strong>of</strong>ten eccentric, rarely 3-septate (7-)9-14(-17)x<br />
(4-)4-5-5-5(-6) /am. Paraphyses numerous, branched and anastomosing, c. l-l-5(-l-7) /xm<br />
wide; apices not swollen or pigmented (green pigment confined to <strong>the</strong> surrounding gel-matrix).<br />
Hypo<strong>the</strong>cium c. 110-200 /u,m tall, hyaline or with very faint olivaceous brown tinge in places.<br />
Fig. 55 Micarea intrusa: a phycobiont cell penetrated by a haustorium <strong>of</strong> an associated mycobiont hypha.<br />
Scale = 10 /u,m.
LICHEN GENUS MICAREA IN EUROPE 139<br />
upper part sometimes pale orange, K+ purple-red (see below); hyphae interwoven, but<br />
becoming vertically orientated towards <strong>the</strong> hymenium, hyaline, c. 1-7-2 fxm wide; ascogenous<br />
hyphae short-celled, to 5 /xm wide, contents sometimes with dilute orange, K+ purple-red<br />
pigment. Excipulum reflexed to below <strong>the</strong> surface <strong>of</strong> surrounding thallus, thin, dilute brownish<br />
or ± hyaline in places, sometimes darkish brown at outer edge; hyphae radiating, branched and<br />
anastomosing, c. 1-5-2 /xm wide.<br />
Pycn/rf/fl not seen.<br />
Chemistry: Thallus surface and sections K— , C-, KC— , PD-,<br />
not analysed by t. I.e.<br />
I-; apo<strong>the</strong>cia sections C—<br />
Observations: This species is placed in Micarea with some hesitation, owing to its unusual<br />
habitat (for a Micarea) and large-celled, thick-walled, non-micareoid phycobiont. However,<br />
much <strong>the</strong> same habitat is shared by M. subnigrata (a species with a typical micareoid phycobiont)<br />
and no o<strong>the</strong>r fungal characters appear to merit its exclusion. It is similar in appearance to M.<br />
subnigrata which can be distinguished by its brown epi<strong>the</strong>cium and usual presence <strong>of</strong> pycnidia<br />
containing helicoid macroconidia. Superficially, M. intrusa is virtually indistinguishable from<br />
Table 5 Species associated with Micarea intrusa in Norway (see text).<br />
Aspicilia morioides<br />
Caloplaca <strong>of</strong>. lamprocheila<br />
Candelariella vitellina<br />
Cornicularia normoerica<br />
Fuscidea cyathoides<br />
F. intercincta<br />
F. tenebrica<br />
Haematomma ochroleucum<br />
Lecanora badia<br />
L. gangaleoides<br />
L. intricata<br />
L. polytropa<br />
L. cf . salina<br />
Lecidea fuliginosa<br />
L. fuscoatra<br />
L. lactea<br />
L. lapicida<br />
L. leucophaea<br />
L. vorticosa<br />
Micarea subnigrata<br />
Opegrapha gyrocarpa<br />
Pertusaria dealbescens<br />
P. lactea<br />
Placopsis gelida<br />
Porina chlorotica<br />
P. guen<strong>the</strong>ri<br />
Ramalina subfarinacea<br />
Rhizocarpon geographicum<br />
R. lecanorinum<br />
R. riparium subsp. lindsayanum<br />
Rinodina gennarii<br />
Schaereria tenebrosa<br />
Stereocaulon vesuvianum<br />
Trapelia involuta<br />
Umbilicaria spodochroa<br />
± vertical<br />
rocks<br />
horizontal<br />
rocks<br />
;
140 BRIAN JOHN COPPINS<br />
forms <strong>of</strong> Scoliciosporum umbrinum with a well developed thallus, but that species has<br />
vermiform or sigmoid-curved spores. The phycobiont in M. intrusa and S. umbrinum appears to<br />
be <strong>the</strong> same; any future critical appraisal <strong>of</strong> <strong>the</strong> delimitation oi Scoliciosporum should seriously<br />
consider <strong>the</strong> generic disposition <strong>of</strong> M. intrusa.<br />
The pale orange, K+ purple-red pigment commonly found in <strong>the</strong> ascogenous hyphae, asci and<br />
spores has not been seen by me in any o<strong>the</strong>r Micarea, although it is conceivable that it is <strong>the</strong><br />
same, or similar, to <strong>the</strong> pigment found in <strong>the</strong> goniocysts <strong>of</strong> M. hedlundii {q. v.).<br />
Habitat and distribution: M. intrusa occurs on hard siliceous (igneous) rocks in communities<br />
referable to <strong>the</strong> Umbilicarietum cylindricae in its broad sense (James etal., 1977). In Nylander's<br />
protologue <strong>of</strong> Lecidea aphanoides it is said to occur on calcareous rock, but this is erroneous;<br />
application <strong>of</strong> 50% HNO3 to fragments <strong>of</strong> substratum from <strong>the</strong> holotype produced no efferves-<br />
cence. The formation <strong>of</strong> small patches amongst o<strong>the</strong>r crustose lichens indicates that M. intrusa<br />
may be, at least facultatively, parasitic. Such suggestions have been previously propounded by<br />
Magnusson (1942), Poelt (1958), and Wirth (1973), all <strong>of</strong> whom related its behaviour to that <strong>of</strong><br />
Lecanora intrudens and Lecidea furvella. In several collections, including <strong>the</strong> type <strong>of</strong> Lecidea<br />
intrusa, it occurs amongst <strong>the</strong> areolae <strong>of</strong> Huilia panaeola, but it is by no means confined to that<br />
'host'.<br />
From her studies on <strong>the</strong> island <strong>of</strong> Sotra in Hordaland, Norway, Miss L. Skjolddal informs me<br />
that M. intrusa occurs on sheltered, sunny, west or south-west facing exposures <strong>of</strong> gneiss and<br />
amphibolite, on steep surfaces or, in one case, on a ± horizontal surface. Miss Skjolddal kindly<br />
provided me with a list <strong>of</strong> associated species (Table 5).<br />
M. intrusa is probably widespread in areas <strong>of</strong> western Europe with exposed, hard, igneous<br />
rocks; however, it is a very inconspicuous species and records are wanting from many potential<br />
localities. I have seen material from sou<strong>the</strong>rn Scandinavia (several localities). North Norway<br />
(Finnmark), and <strong>the</strong> Grampian Mountains <strong>of</strong> Scotland. In addition, Wirth (1973) cites collections<br />
from France (Vosges) and West Germany (Schwarzwald).<br />
Exsiccata: Norrlin & Nyl. Herb. Lich. Fenn. 182 (BM).<br />
18. Micarea leprosula (Th. Fr.) Coppins & A. Fletcher<br />
(Figs 18, 53-54; Map 8)<br />
in Fletcher in Lichenologist 7: 111 (1975). - Bilimbia milliaria y leprosula Th. Fr. , Lich. Scand. 2: 382<br />
(1874). - Micarea violacea var. leprosula (Th. Fr.) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18<br />
(3): 81, 92 (1892). - Bilimbia leprosula (Th. Fr.) H. Olivier in Bull. Geogr. bot. 21: 179 (1911). - Bacidia<br />
leprosula (Th. Fr.) Lettau in Hedwigia 52: 133 (1912). Type: Sweden, Uppsala, Witulfsberg [Wittuls-<br />
berg], 26 vi 1852, Th. M. Fries (UPS - lectotype! [t.l.c. : argopsin and gyrophoric acid]).<br />
Thallus effuse, superficial, <strong>of</strong> convex to subglobose areolae c. 100-350(-400) fxm diam; <strong>the</strong>se<br />
usually coalesce toward <strong>the</strong> centre <strong>of</strong> <strong>the</strong> thallus, whence <strong>the</strong>y may proliferate, producing<br />
secondary granular-areolae, such that <strong>the</strong> thallus becomes thicker (to c. 700 /xm thick). Areolae<br />
blue-grey or, more rarely, grey-brown, matt with a minutely roughened surface with white flecks<br />
(x50 lens); lower sides <strong>of</strong> ± globose areolae, and areolae in shaded situations, <strong>of</strong>ten unpigmented<br />
and greenish white. Areolae fragile and <strong>of</strong>ten breaking down or eroding to form<br />
sorediate patches with irregularly shaped soredial granules c. 20-50 /xm diam. Sections <strong>of</strong> intact<br />
areolae ecorticate and without an amorphous hyaline covering layer; outermost hyphae <strong>of</strong>ten<br />
coloured with greenish or brownish pigment, K— , HNO3-I- reddish. Phycobiont micareoid, cells<br />
4-7 /xm diam.<br />
Apo<strong>the</strong>cia <strong>of</strong>ten absent, at first adnate but <strong>of</strong>ten becoming constricted below, convex to<br />
subglobose, commonly tuberculate, immarginate or rarely very faintly marginate when young,<br />
matt or slightly glossy, dark blue-grey or black, 0- 15-0-4 mm diam, or to 0-7 mm diam when<br />
HNO3-I-<br />
tuberculate. Hymenium 45-60 /xm tall, hyaline or dilute green, but dark green (K— ,<br />
red) in upper part (epi<strong>the</strong>cium). Asci clavate, 40-55x14-16 /xm. Spores oblong-ellipsoid,<br />
oblong-fusiform or clavate-fusiform, <strong>of</strong>ten curved, (l-)3-septate, 14-26(-29)x4-5(-5-5) /xm.<br />
Paraphyses numerous, branched and anastomosing, (0-7-)l-l-5 /xm wide; apices <strong>of</strong>ten more
LICHEN GENUS MICAREA IN EUROPE 141<br />
richly branched, not or only slightly incrassate (to 1-8 /xm wide) and without closely adhering<br />
pigment (epi<strong>the</strong>cial pigment confined to surrounding gel-matrix). Hypo<strong>the</strong>cium c. 35-45 txm<br />
tall, but becoming much taller in markedly convex or tuberculate apo<strong>the</strong>cia, dilute straw- or<br />
fuscous-brown, pigment confined to gel matrix; hyphae interwoven, but becoming outwardly<br />
orientated towards <strong>the</strong> hymenium and excipulum, c. 0-8-1-5 /u,m wide; ascogenous hyphae with<br />
swollen cells, c. 2-6 /am wide. Excipulum distinct in young, moderately convex apo<strong>the</strong>cia, but<br />
reflexed and obscured in subglobose or tuberculate apo<strong>the</strong>cia, dilute fuscous brown (K— );<br />
hyphae radiating, branched and anastomosing, c. 0-5-1 -5 /u,m wide.<br />
Pycnidia not found.<br />
Chemistry: Thallus K-, C+ red, PD-I- red; apo<strong>the</strong>cia in section C-l- red; t.l.c: argopsin and<br />
gyrophoric acid.<br />
Observations: M. leprosula is characterized by its sterile or sparingly fertile, granular thallus<br />
composed <strong>of</strong> blue-grey (or brown-grey), fragile areolae which <strong>of</strong>ten dissolve (or become<br />
abraded) to form contrasting, yellowish green, sorediose patches. These characters are shared<br />
by <strong>the</strong> much rarer M. subleprosula, and <strong>the</strong> thalli <strong>of</strong> both react C+ red. However, M.<br />
subleprosula can be separated by <strong>the</strong> PD-I- yellow reaction <strong>of</strong> its thallus due to <strong>the</strong> presence <strong>of</strong><br />
alectorialic acid. When fertile M. subleprosula is found to have much larger, mostly 7-septate<br />
spores. M. leprosula is usually distinctive in <strong>the</strong> field (although material should always be<br />
checked with M. subleprosula in mind) but can easily be overlooked when it occurs on<br />
bryophytes on shaded rocks or trees, whence <strong>the</strong> thallus <strong>of</strong>ten lacks its characteristic bluish tinge<br />
and <strong>the</strong> areolae are thinly scattered. The species is usually sterile but fertile specimens are<br />
occasionally encountered, especially in sheltered (but not deeply shaded) situations, such as on<br />
boulders and old walls in woodland.<br />
Hedlund (1892) regarded M. leprosula as a variety <strong>of</strong> M. peliocarpa (q.v.) and <strong>the</strong> apo<strong>the</strong>cia<br />
<strong>of</strong> <strong>the</strong> two species are somewhat similar. However, those <strong>of</strong> M. leprosula exhibit several minor,<br />
yet significant, differences; spores tend to be slightly longer; apo<strong>the</strong>cia tend to be more<br />
markedly convex and more frequently tuberculate; hypo<strong>the</strong>cium and excipulum are dilutely<br />
pigmented with a dull brown pigment; and paraphyses never become thickened and pigmented<br />
at <strong>the</strong>ir apices. The two species also differ in several fundamental aspects <strong>of</strong> thallus morphology<br />
and chemistry. With regard to <strong>the</strong> production <strong>of</strong> argopsin and to <strong>the</strong> presence <strong>of</strong> a dull brown<br />
pigment in <strong>the</strong> excipulum and hypo<strong>the</strong>cium M. leprosula shows some affinity to M. lignaria var.<br />
lignaria, with which it <strong>of</strong>ten occurs. However, M. lignaria lacks gyrophoric acid, has usually<br />
longer and more septate spores, and somewhat stouter and only sparingly branched paraphyses.<br />
The areolae <strong>of</strong> M. leprosula and M. subleprosula differ from those <strong>of</strong> such species as M. lignaria,<br />
M. peliocarpa, and M. cinerea, in lacking an amorphous hyaline covering layer (see 'morpho-<br />
logy')-<br />
Habitat and distribution: M. leprosula is widely distributed in upland areas in <strong>the</strong> north and<br />
west <strong>of</strong> Britain, occurring at altitudes from about sea-level to at least 1100 m. It is found, usually<br />
on moribund bryophytes (e.g. Andraea spp. and Rhacomitrium spp.) on rocky ledges, boulders,<br />
and old walls, in exposed hilly areas, mountain sides or woodlands. Associated lichens noted<br />
amongst <strong>British</strong> collections include Arthrorhaphis citrinella, Baeomyces rufus, Belonia incar-<br />
nata, Cladonia cervicornis, C. coccifera, C. crispata, C. portentosa, C. squamosa, C. subcervicornis,<br />
Coelocaulon aculeatum, Lecidea granulosa, L. icmalea, Lepraria neglecta, Micarea<br />
lignaria, M. peliocarpa, and Vorarlbergia renitens. From Scotland I have seen two corticolous<br />
specimens, on trunks <strong>of</strong> Alnus and Betula; associated species present included Cladonia<br />
coniocraea, C. squamosa, Lecidea icmalea, Micarea melaena, and Platismatia glauca.<br />
There are two outlying localities in <strong>the</strong> lowlands <strong>of</strong> sou<strong>the</strong>rn England. In Dorset it occurs on<br />
waste heaps <strong>of</strong> slag clay with M. lignaria (q. v.); and in Kent it was found with M. peliocarpa on<br />
wood chips lying at <strong>the</strong> side <strong>of</strong> a woodland track in a chestnut (Castanea) coppice.<br />
M. leprosula is poorly recorded outside Britain, probably because it is so <strong>of</strong>ten sterile. To<br />
date, I have seen one specimen from Norway (Hordaland) and several collections from mid- and<br />
sou<strong>the</strong>rn Sweden. Fur<strong>the</strong>r south it seems to be well represented in <strong>the</strong> Alps, and I have seen one
142 BRIAN JOHN COPPINS<br />
Map 8 Micarea leprosula # 1950 onwards O Before 1950<br />
collection from Bohemia (Czechoslovakia). This Bohemian record, and ano<strong>the</strong>r from <strong>the</strong> Black<br />
Forest (Schwarzwald) were made from exsiccate specimens {Kutdk 417 and Migula 1) <strong>of</strong> M.<br />
lignaria growing on wooden ro<strong>of</strong> shingles.<br />
Exsiccata: Kutak Lich. Bohem. 417 p. min. p. (0). Magnusson Lich. Sel. Scand. 208 (BM, C, GZU).<br />
Migula Crypt. Germ. \p. min. p. (C, E, M).<br />
19a. Micarea lignaria (Ach.) Hedl. var. lignaria<br />
(Figs lA, 3D-E, 19, 45, 53-54; Map 9)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 82, 93 (1892). - Lecidea lignaria Ach. in K. svenska<br />
VetenskAkad. Handl. 1808: 236 (1808). - Bilimbia lignaria (Ach.) Massal. , Ric. Lich. Crost. : 121 (1852).<br />
- Bacidia lignaria (Ach.) Lettau in Hedwigia 52: 132 (1912). Type: Sweden: 'Svecia', on lignum,<br />
(H-ACH 265 - lectotype; BM - ? isotype!).<br />
Lecidea milliaria Fr. in K. svenska VetenskAkad. Handl. 1822: 255 (1822). Type: Sweden, Smaland,<br />
Femsjo, on lignum, E. M. Fries (UPS - lectotype! [t.l.c: argopsin]).<br />
Lecidea geomaea Taylor in Mackay, Fl. Hib. 2: VIA (1836). Type: Ireland, South Kerry, Dunkerron, T.<br />
Taylor (BM - lectotype!; BM ex K - isolectotype!).<br />
Bilimbia milliaria a. [var.] lignaria** [f.] calamophila Korber, Parerga lich. : 171 (1860). Type: Germany:<br />
'an Schilfrohrdachern um Miinster', J. G. F. X. Lahm (L 910, 144-1685 - lectotype!; L 910, 144-1689 -<br />
isolectotype! [t.l.c: argopsin].<br />
Bilimbia milliaria (3. [var.] saxicola Korber, Parerga lich. : 171 (1860). Type: Germany: 'Bilimbia trochiscus<br />
Korb. prim, B. miliaria p. saxicola Kb.! Extersteine', on sandstone, ? Beckhaus (WRSL- lectotype!).
LICHEN GENUS MICAREA IN EUROPE 143<br />
Paratype: Germany: 'Ibbenbiiren in Westfalen, Lahm, Hb. Hepp.', on sandstone (BM!).<br />
Stereocauliscum gomphillaceum Nyl. in Flora, Jena, 48: 211 (1865). - Micarea gomphillacea (Nyl.) Vezda<br />
in Folia geobot. phytotax., Praha 5: 321 (1970). - Bilimbia milliaria f. [as 'f?'] gomphillacea (Nyl.)<br />
Blomb. & Forss., Enum. PI. Scand.: 82 (1880). - Micarea lignaria [as 'ligniaria'] f. gomphillacea (Nyl.)<br />
Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 94 (1892). Type: Finland, Tavastia australis,<br />
HoUola, Tiirismaa, Pirunpesa, [on granite rock], 1863,/. P. Norrlin (H-NYL 40217 -lectotype! [t.l.c:<br />
argopsin];isolectotypes: H-NYL 40214!, 402151,40216!, 40218! and p.m. 631!).<br />
Lecidea sabuletorum var. milliaria f. nigrata Nyl. in Not. Sdllsk. Fauna Fl. fenn. Forh. 8: 151 (1866). Type:<br />
USSR, Kola Peninsular, Lapponiaponojensis,Ponoi, 1863, M /. Fe/Zman (H-holotype!;H-isotype!).<br />
See note (i) below.<br />
Lecidea milliaria var. triseptata Nyl. in Lamy in Bull. Soc. bot. Fr. 25: 441 (1878). - Bilimbia triseptata<br />
(Nyl.) Arnold in Flora, Jena 67: 572 (1884), non Bacidia triseptata (Hepp in ZoU.) Zahlbr. Type: France:<br />
Haute Vienne, 'Sur des roches entre Chalard et St Junien, Lamy (H-NYL 18367! - lectotype, as 'L.<br />
milliaria var. 3-septata Nyl.'). See note (ii) below.<br />
Lecidea meizospora Harm, in Bull. Seanc. Soc. Sci. Nancy II, 33 (1898): 63 (1899). - Bacidia meizospora<br />
(Harm.) Zahlbr., Cat. lich. univ. 4: 122 (1926). Type: France, Vosges, Docelles, on Calluna roots, V.<br />
and H. Claudel & F. J. Harmand (ANGUC - lectotype!).<br />
Lecidea trisepta var. polytropoides Vainio in Ark. Bot. 8 (4): 106 (1909). Type: USSR, Magadanskya<br />
Oblast (N.E. Siberia), 'Pitlekai, peninsula Jinretlen' [c. 67°7'N 173°24'W], on plant debris, 1878-9, E.<br />
Almquist (TUR-VAINIO 21274, fertile part- lectotype!). See note (iii) below.<br />
Patellaria milliaria var. xylophila Wallr., Fl. crypt. Germ. 1: 349 (1831); nom. inval. (Art. 26).<br />
Lecidea milliaria var. saxigena Leighton, Lich. Brit. Exs. 210 (1856); nom. nudum (Art. 32); see note<br />
under M. peliocarpa. Type: England, Shropshire, Neescliff [Nesscliff] Hill, Leighton, Lich. Brit. Exs.<br />
210 (E- lectotype!; isolectotypes: BM! DBN! MANCH! NMW!).<br />
Notes: (i) The type material mainly consists <strong>of</strong> <strong>the</strong> black stroma <strong>of</strong> Bryomyces gymnomitrii Dobb. on<br />
Gymnomitrium sp. , with only a few apo<strong>the</strong>cia <strong>of</strong> M. lignaria present.<br />
(ii) Lamy (loc. cit.) gives <strong>the</strong> locality as 'entre le Chalard et Saint-Yrieix'. The label on <strong>the</strong> lectotype<br />
specimen is entirely in Nylander's handwriting and <strong>the</strong>re may have been an error in transcription.<br />
(iii) Vainio's diagnosis included <strong>the</strong> description <strong>of</strong> soralia. A small portion <strong>of</strong> <strong>the</strong> type specimen is a<br />
sterile sorediate thallus (C- , PD-<br />
phycobiont cells 7-14 /am diam) but it is not a Micarea and is not part <strong>of</strong><br />
;<br />
<strong>the</strong> larger, fertile portion. The latter belongs to Micarea lignaria and is chosen as lectotype.<br />
Thallus effuse, partially endoxylic or endocuticular, but more usually developed superficially<br />
as convex to ± globose areolae. Areolae c. 80-250(-300) /xm diam, discrete, scattered to<br />
contiguous, sometimes coalescing to form an uneven crust, grey-white and ± glossy to<br />
grey-green or bluish grey and matt, sometimes brown-grey due to <strong>the</strong> ramification through <strong>the</strong>ir<br />
surface <strong>of</strong> brown, toruloid hyphae (cells c. 5-7x4-5-7 /xm) belonging to an unknown fungus.<br />
Areolae in section, with a hyaline amorphous covering layer (c. 4—12 /xm tall); walls <strong>of</strong> hyphae in<br />
outermost (exposed) parts <strong>of</strong>ten blue-green (K-, HNO3+ red); algal layer usually in direct<br />
contact with substratum, but in larger areolae <strong>the</strong> algal layer may be c. 90-135 ixva tall lying<br />
above a medulla. Medulla, when present, up to c. 100 ^im tall, consisting <strong>of</strong> loosely interwoven<br />
hyphae c. 1-1-5 /xm wide, intermixed with dead algal cells and debris derived from <strong>the</strong><br />
substratum. Phycobiont mic2iVQO\d, cells c. A-1 /xm diam.<br />
Apo<strong>the</strong>cia scattered, more usually numerous and crowded, <strong>of</strong>ten confluent, immarginate,<br />
convex-hemispherical, soon becoming constricted at <strong>the</strong> base and hence ± globose, occasionally<br />
becoming stipitate with a grey-black 'stipe' up to 1 mm tall ('f. gomphillacea''), black, matt or<br />
slightly glossy, rarely blue-grey (shade forms), 0-15-0-6(-0-9) mm diam. Hymenium 50-75 /xm,<br />
<strong>of</strong>ten not sharply delimited from <strong>the</strong> hypo<strong>the</strong>cium, dilute olivaceous or aeruginose-green (K-<br />
HNO3-I- red) in upper, and sometimes lower, part(s) (<strong>the</strong> middle part <strong>of</strong>ten being ± hyaline); in<br />
<strong>the</strong> uppermost part (epi<strong>the</strong>cium) <strong>the</strong> pigment is more dense and is present not only in <strong>the</strong> gel<br />
matrix but also in <strong>the</strong> walls <strong>of</strong> <strong>the</strong> paraphyses; in reflexed parts <strong>of</strong> <strong>the</strong> hymenium towards <strong>the</strong> base<br />
<strong>of</strong> <strong>the</strong> apo<strong>the</strong>cium <strong>the</strong> green pigment is <strong>of</strong>ten replaced by a dilute brown pigment; minute (less<br />
than 2x1 yu,m) granules <strong>of</strong> violet (K+ intense aeruginose) pigment are sometimes present and<br />
lightly scattered through <strong>the</strong> hymenium. Asci clavate, 45-50x11-19 /xm. Spores fusiform,<br />
straight or slightly curved, (0-)3-7 septate, 16-36(-38)x4-6(-7) /xm. Paraphyses numerous,<br />
simple or slightly branched above, but more richly branched in <strong>the</strong> transition zone between <strong>the</strong><br />
hymenium and excipulum, sometimes anastomosing, 1-3-1-8 /xm wide; apices <strong>of</strong>ten slightly<br />
,
144 BRIAN JOHN COPPINS<br />
incrassate and coloured with green or greenish brown pigment and <strong>the</strong>n up to 3 /xm wide, usually<br />
coherent (even in K) and, toge<strong>the</strong>r with <strong>the</strong> surrounding pigmented matrix, form an epi<strong>the</strong>cium.<br />
Hypo<strong>the</strong>cium c. 100-230(-350) /xm tall, dilutely pigmented with pigment confined to gel matrix;<br />
upper part dull aeruginose or olive-brownish K- or + dulling, HNO3+ reddish); lower part<br />
<strong>of</strong>ten without greenish tinge, and <strong>the</strong>n being dilute brownish or ± hyaline; hyphae interwoven or<br />
some ± vertically orientated near <strong>the</strong> hymenium, c. 1-1-7 /am wide; ascogenous hyphae with<br />
short, swollen cells c. 2-5-4 /xm wide. Excipulum ± distinct in sections <strong>of</strong> young, hemispherical<br />
apo<strong>the</strong>cia, but soon becoming strongly reflexed and not sharply delimited from <strong>the</strong> hymenium,<br />
dilute brown or sometimes darkish brown along <strong>the</strong> outer edge. Hypo<strong>the</strong>cial and excipular<br />
tissues sometimes elongated vertically to form a stipe ('f. gomphillaced')<br />
Pycnidia inconspicuous, ± immersed, with walls green (K-, HNO3-I- red) in exposed upper<br />
parts and ± hyaline in immersed lower parts; <strong>of</strong> three types [only type (c) is common]: (a) c. 100<br />
ixm diam; conidia {macroconidia) curved or hamate, 0-3-septate, 16-22 Xc. 1 /xm; (b) c. 100-140<br />
/xm diam; conidia (mesoconidia) ± cylindrical, oblong-ellipsoid, obovoid or oblong-obovoid,<br />
usually distinctly truncate at proximal end, <strong>of</strong>ten 2-3-guttulate, 4-7(-7-6)xl-2-l-8 fxm; (c) c.<br />
40-50 ^im diam; conidia (microconidia) narrowly cylindrical, only faintly truncated at proximal<br />
end, eguttulate, (4-5-)5-7(-8)x 0-8-1 /xm.<br />
Chemistry: Thallus K-, C-, KC-, PD-I- red; sections <strong>of</strong> apo<strong>the</strong>cia C-, PD-; t.l.c:<br />
argopsin.<br />
Observations: Micarea lignaria is characterized by its whitish to grey, convex to ± globose<br />
areolae which are PD+ red (argopsin; but see var. endoleuca), black, markedly convex to ±<br />
globose apo<strong>the</strong>cia, green upper hymenium, dilute greenish or dilute olive-brownish hypo<strong>the</strong>cium,<br />
'^nd 3-7-septate, fusiform spores. When on lignum <strong>the</strong> thallus is <strong>of</strong>ten reduced to small<br />
scattered areolae and sometimes it is ± entirely endoxylic. The apo<strong>the</strong>cia are less variable in<br />
appearance, and pale (shade) forms are very rare. However, <strong>the</strong> apo<strong>the</strong>cia are occasionally<br />
found (especially on rock in dry underhangs) to be stipitate with a 'stipe' (composed <strong>of</strong> vertically<br />
extended hypo<strong>the</strong>cial and excipular tissues) up to 0-4 mm, or even 1 mm, tall (Fig. 3E). These<br />
forms have been ascribed <strong>the</strong> status <strong>of</strong> form, variety, species and even genus, viz.: Nylander's<br />
new species and genus Stereocauliscum gomphillaceum. In <strong>the</strong> type material <strong>of</strong> this, some<br />
'stipes' are extremely tall (up to 1 mm) and composed <strong>of</strong> vertically proliferating apo<strong>the</strong>cia, with<br />
<strong>the</strong> apical apo<strong>the</strong>cia being <strong>the</strong> youngest; <strong>the</strong> apo<strong>the</strong>cia are immature (or arrested in <strong>the</strong>ir<br />
development) with few asci and spores. In my opinion <strong>the</strong>se stalked forms <strong>of</strong> A/, lignaria result<br />
from abnormal development in response to adverse environmental conditions and are, <strong>the</strong>re-<br />
fore, not worthy <strong>of</strong> taxonomic recognition at any rank.<br />
M. lignaria is <strong>of</strong>ten confused with M. cinerea and M. peliocarpa, but <strong>the</strong>se two species have<br />
usually more flattened and <strong>of</strong>ten paler apo<strong>the</strong>cia, more richly branched paraphyses, and a<br />
hyaline hypo<strong>the</strong>cium. In addition, <strong>the</strong>y contain gyrophoric acid, resulting in <strong>the</strong> C-l- orange-red<br />
(quickly fading) , PD<br />
— reactions <strong>of</strong> <strong>the</strong>ir thalli and apo<strong>the</strong>cia. M. ternaria {q. v. ) is very similar to<br />
M. lignaria, but has more flattened apo<strong>the</strong>cia with a more discernible excipulum and spores<br />
which are never more than 3-septate; fur<strong>the</strong>rmore, it lacks any lichen substances.<br />
Habitat and distribution: M. lignaria occurs on a wide range <strong>of</strong> substrata, but is most common<br />
in upland districts, growing over bryophytes or plant debris on siliceous rocks and walls, or on<br />
exposed peaty ground. Associated lichens found in such habitats in Britain include<br />
Arthrorhaphis citrinella, Baeomyces rufus, Cladonia coccifera, C. floerkeana, C. squamosa, C.<br />
subcervicornis, Coelocaulon aculeatum s. lat., Lecidea granulosa, L. icmalea, Lecidoma demissum,<br />
Lepraria neglecta, Micarea leprosula, M. peliocarpa, Ochrolechia frigida, Parmelia<br />
saxatilis, and Pseudephebe pubescens. It <strong>of</strong>ten grows over bryophyte mats that are heavily<br />
invaded by gelatinous algae, and in such situations in Scotland it has been found with <strong>the</strong> rare, or<br />
overlooked, Arctomia delicatula and Belonia incarnata. When growing directly on rock M.<br />
lignaria is mostly confined to sheltered, shaded situations and is sometimes present in <strong>the</strong><br />
Micareetum sylvicolae in rock underhangs. It is usually present in abundance in <strong>the</strong> old lead and<br />
zinc mine workings <strong>of</strong> <strong>the</strong> Pennines and Scotland, where it grows over decaying bryophytes and<br />
.
LICHEN GENUS MICAREA IN EUROPE 145<br />
Map 9 Micarea lignaria var. lignaria # 1950 onwards O Before 1950<br />
plant debris, loose stones, pieces <strong>of</strong> timber and sack-cloth. Mr V. Giavarini has recently found it<br />
in Dorset, growing on waste heaps <strong>of</strong> slag clay, in <strong>the</strong> company <strong>of</strong> M. leprosula, Baeomyces<br />
roseus, Cladonia arbuscula, C. ciliata, C. furcata, C. portentosa, and Coeldcaulon aculeatum s.<br />
lat. M. lignaria is very rare in <strong>the</strong> lowlands <strong>of</strong> south-east England, but has been found <strong>the</strong>re<br />
growing directly on <strong>the</strong> rock <strong>of</strong> east or north facing sandstone walls and churchyard memorials,<br />
and also on natural sandstone outcrops <strong>of</strong> <strong>the</strong> Sussex Weald.<br />
In upland areas M. lignaria is frequently found on <strong>the</strong> exposed lignum <strong>of</strong> fallen trunks<br />
(especially <strong>of</strong> conifers) and old timberwork, with, for example, Cladonia spp., Hypogymnia<br />
physodes, Micarea peliocarpa, Mycoblastus sterilis, Ochrolechia turneri, Parmelia saxatilis, and<br />
Lecanora polytropa (worked timber) . Occurrences <strong>of</strong> M. lignaria on <strong>the</strong> bark <strong>of</strong> healthy trees<br />
are ra<strong>the</strong>r rare, and in Britain are confined to <strong>the</strong> high rainfall areas <strong>of</strong> <strong>the</strong> north and west, where<br />
it has been collected on Alnus, Betula, Fraxinus, Ilex, Quercus, and old Sambucus.<br />
Reports <strong>of</strong> M. lignaria on mosses on limestone rocks usually result from <strong>the</strong> misidentification<br />
<strong>of</strong> Bacidia sabuletorum or Toninia lobulata. However, on a few occasions I have seen M. lignaria<br />
growing on thick bryophyte cushions or mats at high altitudes over limestone in <strong>the</strong> north<br />
Pennines, and over calcareous mica-schist in <strong>the</strong> Breadalbane Mountains; in such situations <strong>the</strong><br />
pH and calcium content <strong>of</strong> <strong>the</strong> substratum is presumably kept low by <strong>the</strong> leaching effect <strong>of</strong> <strong>the</strong>'<br />
high rainfall in those areas.<br />
The altitude range <strong>of</strong> M. lignaria in Britain is from sea-level to about 1200 m (Ben Lawers),<br />
although it may well occur at higher altitudes in <strong>the</strong> Ben Nevis group and <strong>the</strong> Cairngorni<br />
Mountains. Higher altitudes are attained in <strong>the</strong> mountains <strong>of</strong> central Europe, from where it has
146<br />
BRIAN JOHN COPPINS<br />
been collected at 2800 m in <strong>the</strong> Austrian Tirol, c. 2000 m in <strong>the</strong> Tatry mountains <strong>of</strong> Poland and<br />
Czechoslovakia, and at nearly 2500 m in <strong>the</strong> Transylvanian Alps <strong>of</strong> Romania.<br />
M. Ugnaria is common in nor<strong>the</strong>rn and western Britain, but is very rare, with only a few<br />
scattered records, in lowland, sou<strong>the</strong>rn England east <strong>of</strong> Devon. It is widely distributed<br />
throughout most <strong>of</strong> Europe, although in sou<strong>the</strong>rn Europe it appears to be confined to <strong>the</strong><br />
mountains: like most species <strong>of</strong> Micarea it seems to avoid <strong>the</strong> lowland Mediterranean regions.<br />
Records suggest that it is rare in arctic Fennoscandia, although I have seen material from north<br />
Finland and <strong>the</strong> Kola Peninsula. It has been found in <strong>the</strong> Azores at altitudes <strong>of</strong> about 900 m but I<br />
do not know <strong>of</strong> it from elsewhere in Macaronesia. From outside Europe I have seen material<br />
from <strong>the</strong> north coast <strong>of</strong> western Siberia, <strong>the</strong> Franconia Mountains (at c. 1450 m) <strong>of</strong> New<br />
Hampshire in <strong>the</strong> USA, and from Brazil (Serra Montiqueira, at c. 1900 m).<br />
Exsiccata: Anzi Lich. Lang. 148 (BM). Arnold Lich. Exs. 348A, B (BM ex K, L, M). Bohler Lich. Brit.<br />
85 (E). Claudel & Harm. Lich. Gall. 43 (BM). Cumm. Dec. N. Am. Lich. ed. I: 302 (BM, C), ed. 2: 232<br />
(M). Flotow Lich. Exs. 129A, E and 131 (UPS). Fries Lich. Suec. 29 (E, M, UPS); 98 (E). Harm. Guide 91<br />
(UPS). Harm. Lich. Loth. 852 (M). Hav. Lich. Norv. 555 (C). Hertel Lecid. 34 (E, GZU). Johnson Lich.<br />
Herb. 453 (E). Kalb Lich. neotropici 22, 186 (hb Kalb). Krypt. Exs. Vindob. 658 (BM, BM ex K, C, M).<br />
Kutak Lich. Bohem. 417 (O). Larb. Lich. Caes. Sarg. 83 (BM). Larb. Lich. Herb. 272 (BM). Leighton<br />
Lich. Brit. 210 (BM, DBN, E, MANCH, NMW), 238 p.p. (BON, DBN, E, M, MANCH), 386 (BM,<br />
BON, DBN, E, FRS, M). Lojka Lich. Hung. 61 (BM, BM ex K, M). Malme Lich. Suec. 288 (C, M, S).<br />
Migula Crypt. Germ. Ip.p. (BM, C, M, MANCH); 226 (BM, C, MANCH). Moug. & Nestl. Stirp. Crypt.<br />
1430 (E). Mudd Lich. Brit. 156 (BM, E, M, MANCH), 157 (E, MANCH), 158 (BM, E, M, MANCH).<br />
Norrlin & Nyl. Herb. Lich. Fenn. 319A, B (BM, C, H). Oliv. Lich. Orne 344 (M, S). Pisut Lich. Slov. 156<br />
(BM, M). Rabenh. Lich. Eur. 322, 603 (BM, M). Roum. Lich. Gall. 193 (BM), 232 (BM, M). Samp. Lich.<br />
Port. 147 (LD). Schaerer Lich. Helv. 196p.p. (BM, BM ex K, E). Vezda Lich. Bohem. 133, 258 (LD, M).<br />
Vezda Lich. Sel. 516, 858, 1036, 1088 (BM). Zwackh. Lich. Exs. Ill (BM).<br />
19b. Micarea lignaria var. endoleuca (Leighton) Coppins, comb. nov.<br />
(Figs 53-54; Map 10)<br />
Lecidea milliaria var. endoleuca Leighton, Lich. Fl. Brit., edn 3: 363 (1879). Type: Ireland, West Galway,<br />
on<strong>the</strong>Doughraugh, 1875, C. Lar/?fl/e5^ieA-(BMexK-lectotype!; BM-? isolectotype!).<br />
Thallus identical to var. lignaria except that <strong>the</strong> areolae usually have dull yellowish<br />
(isabelline) tinge.<br />
Apo<strong>the</strong>cia and pycnidia (containing microconidia) identical to var. lignaria; pycnidia containing<br />
macroconidia or mesoconidia not known.<br />
Chemistry: Thallus K- or Kf+ yellow, C+ yellow-orange (persistent), KC-I- reddish orange<br />
(persistent), PD-; sections <strong>of</strong> apo<strong>the</strong>cia C-; t.l.c: xanthone(s), possibly artho<strong>the</strong>lin and<br />
thiophaninic acid.<br />
Observations (including habitat and distribution) : My<br />
preliminary studies on M. lignaria found<br />
most specimens to have a C-, PD+ red thallus reaction, but a few o<strong>the</strong>rwise ± identical<br />
specimens from western Ireland, north Wales, and western Scotland gave C+ persistent orange,<br />
PD- reactions. Subsequent t.l.c. analysis proved <strong>the</strong> normal form to have <strong>the</strong> <strong>the</strong>n unidentified,<br />
P+ red compound found also in Phyllopsora rosei (Coppins & James, 1979), and <strong>the</strong> anomalous<br />
form to contain two xanthones (possibly artho<strong>the</strong>lin and thiophaninic acid). Xanthones <strong>of</strong>ten<br />
impart a yellowish tinge to lichen thalli (e.g. Lecidella elaeochroma, L. scabra, Pertusaria<br />
flavida, and P. hymenea) and this is <strong>the</strong> case for <strong>the</strong> xanthone containing race <strong>of</strong> M. lignaria; <strong>the</strong><br />
colour difference is most evident when <strong>the</strong> two races occur toge<strong>the</strong>r. Of all <strong>the</strong> types <strong>of</strong> names<br />
attributable to M. lignaria s. ampL, only that applicable to <strong>the</strong> 'var. endoleuca Leighton' was<br />
found to contain xanthones, and that name is here adopted at <strong>the</strong> same rank. Although <strong>the</strong> var.<br />
endoleuca is morphologically identical to <strong>the</strong> var. lignaria it has a sympatric but much more<br />
restricted geographical range. In <strong>the</strong> <strong>British</strong> Isles it is confined to <strong>the</strong> more oceanic districts with<br />
one ra<strong>the</strong>r anomalous occurrence in Surrey (Leith Hill). Fur<strong>the</strong>rmore, it appears to have a lower<br />
altitudinal range, with a collection at 600 m in Snowdonia representing its known upper limit in
LICHEN GENUS MICAREA IN EUROPE 147<br />
Map 10 Micarea lignaria var. endoleuca # 1950 onwards O Before 1950<br />
<strong>the</strong> <strong>British</strong> Isles. Only four examples <strong>of</strong> var. endoleuca have been discovered from outside<br />
Britain: from Baden (Heidelberg) and Bavaria (Bayreuth) in Germany, Bern (Langnau) in<br />
Switzerland, and Trentino (Appiano) in nor<strong>the</strong>rn Italy. It should be checked for amongst<br />
material from o<strong>the</strong>r regions, especially south-west Norway, Bretagne (France) and <strong>the</strong> Pyre-<br />
nees.<br />
I believe that <strong>the</strong> var. endoleuca should be retained at varietal rank. However, as <strong>the</strong><br />
depsidone argopsin (I'-chloropannarin) <strong>of</strong> var. lignaria is probably biogenetically distinct from<br />
<strong>the</strong> xanthones <strong>of</strong> var. endoleuca, a case could be made for its recognition at species rank. A<br />
comparative example is Parmeliopsis ambigua versus P. hyperopia (see Hawksworth, 1976).<br />
20. Micarea lithinella (Ny 1 ) Hedl<br />
.<br />
(Figs4A,20A-B;Mapll)<br />
.<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 97 (1892). - Lecidea lithinella Nyl. in Flora, Jena<br />
63: 390 (1880). - Lecidea lithinella Nyl. in Flora, Jena 45: 464 (1862); nom. nudum (Art. 32). Type:<br />
Germany, Bayern, 'Sandsteineblocke am Waldwege von Banz nach Altenbanz in Oberfranken', ix<br />
1860, F. Arnold 957 (M - lectotype!; H-NYL 19192 - isolectotype (fragment)!).<br />
Thallus effuse, saxicolous, <strong>of</strong>ten a thin indistinct crust between protruding grains <strong>of</strong> rock, but<br />
somes developing irregularly rounded, convex, more rarely subglobose, whitish areolae, c.<br />
40-100 /xm diam. Phycobiont micareoid, cells 4—6-5 /xm diam.<br />
Apo<strong>the</strong>cia numerous, immarginate, adnate, convex, apparently never tuberculate, pallid or
148 BRIAN JOHN COPPINS<br />
dull yellow-orange to reddish brown (never black), (O-l-)O- 15-0-4 mm diam. Hymenium 35-50<br />
^im tall, hyaline but <strong>of</strong>ten tinged dilute straw in upper part. Asci cylindrical-elavate, 35-50x8-13<br />
/xm. Spores ovoid or fusiform-ellipsoid, simple, 6-5-9-5X 2-8-4 ^tm. Paraphyses scanty, simple<br />
or irregularly forked above, 0-7-1 ^tm wide but sometimes gradually widening towards <strong>the</strong><br />
apices and up to 1-5 //-m wide, hyaline throughout. Hypo<strong>the</strong>cium c. 50-70 fxm tall, yellow-straw<br />
to dilute orange-brown, K- , HNO3-<br />
pigment confined to gel-matrix , and hyphal walls hyahne.<br />
Excipulum absent (not detected even in sections <strong>of</strong> young apo<strong>the</strong>cia).<br />
Pycnidia apparently rare, very inconspicuous, immersed, c. 40 /am diam, with hyaline walls.<br />
Conidia (microconidia) narrowly cylindrical, 4-5-7x0-8-1 fim.<br />
Chemistry: Thallus C— , K—<br />
, PD-; section <strong>of</strong> apo<strong>the</strong>cia C-; t.l.c: not tested.<br />
Observations: M. lithinella exhibits little variation except that between, and sometimes<br />
within, collections <strong>the</strong>re is an intergradation from pallid apo<strong>the</strong>cia with a dull yellowish<br />
hypo<strong>the</strong>cium to darker apo<strong>the</strong>cia with an orange-brown hypo<strong>the</strong>cium. This variation is correlated<br />
with differences in exposure to light. M. lithinella is easily confused with forms <strong>of</strong> M.<br />
bauschiana with pallid apo<strong>the</strong>cia. Difficult specimens can be determined by a careful examination<br />
<strong>of</strong> <strong>the</strong> phycobiont, which is micareoid in M. lithinella but non-micareoid (cells c. 7-13 jxm<br />
diam) in M. bauschiana. M. lutulata has darker apo<strong>the</strong>cia which are commonly tuberculate, a<br />
darker (opaque) hypo<strong>the</strong>cium, an <strong>of</strong>ten greenish hymenium, and non-micareoid phycobiont.<br />
M. muhrii is normally lignicolous but has once been found on rocks; it has adnate apo<strong>the</strong>cia like<br />
M. lithinella but <strong>the</strong>y are usually darker and have a taller, dark reddish brown hypo<strong>the</strong>cium, and<br />
usually a greenish upper hymenium. M. myriocarpa shares a similarly coloured hypo<strong>the</strong>cium<br />
with M. lithinella, but has smaller, globose to tuberculate apo<strong>the</strong>cia and smaller, 1-septate<br />
spores.<br />
In <strong>the</strong>ir keys to Micarea Vezda & Wirth (1976) and Poelt & Vezda (1977) state that M.<br />
lithinella has a K+ violet hymenium and <strong>of</strong>ten 1-septate spores; this is clearly an error, probably<br />
due to confusion with saxicolous forms <strong>of</strong> M. denigrata or M. prasina.<br />
Habitat and distribution: M. lithinella occurs on acidic rocks, especially hard sandstone.<br />
Associated species encountered on <strong>the</strong> specimens examined include Baeomyces rufus (parasi-<br />
tized by Thelocarpon lichenicola on <strong>the</strong> lectotype <strong>of</strong> Lecidea lithinella), Huilia crustulata<br />
Rhizocarpon obscuratum, Scoliciosporum umbrinum, Trapelia coarctata, and T. aff . obtegens.<br />
This list <strong>of</strong> associates, plus information on some <strong>of</strong> <strong>the</strong> packets, suggest that M. lithinella has<br />
mostly been found in humid situations on outcrops and boulders by woodland roads. It was<br />
collected by Arnold, Lahm and von Zwackh from several areas <strong>of</strong> Germany: Bayern, Nordrhein-Westfalen<br />
and Baden-Wiirttemberg, respectively. In Sweden it was collected in Uppland<br />
by Hedlund who (1892: 97) also cited a collection by Blomberg from Sodermanland. One <strong>British</strong><br />
specimen has recently been found: South-east York (V.C. 61), Wharram Quarry Nature<br />
Reserve, 44/85. 65, 1969, Coppins (E). At this locality M. lithinella was growing with Scoliciosporum<br />
umbrinum on <strong>the</strong> underside <strong>of</strong> a large flint; <strong>the</strong> upper, more exposed, part <strong>of</strong> <strong>the</strong> flint<br />
was colonized by Lecidea erratica and Rhizocarpon obscuratum.<br />
Exsiccata: Arnold Lick. Exs. 836 (BM ex K, H-NYL 19190, M). Lojka Lick. Univ. 233 (M). Malme<br />
Lick. Suec. 125 (M, S). Zwackh Lich. Exs. 590 (H-NYL p.m. 5404, M).<br />
21. Micarea lutulata (Nyl.) Coppins<br />
(Figs 20c, 44B; Map 11)<br />
in D. Hawksw., P. James, & Coppins in Lichenologist 12: 107 (1980). - Lecidea lutulata Nyl. in Flora,<br />
Jena56: 297 (1873). Type: Jersey, Rozel meadow, bases <strong>of</strong> rocks, 1873, C. Larbalestier (H-NYL 10696lectotype!;<br />
BM-isolectotype!; M- probable isolectotype!). See note below.<br />
Lecidea laxulaNyl. in Flora, Jena 5S: 11 (1875). Type: Finland, Tavastiaaustraiis,Luhanka, Hietala, 1874,<br />
E. A. Lang [Vainio] 303 (H-NYL 20689 - lectotype!; H - isolectotype!).<br />
Lecidea poliodes Nyl. in Flora, Jena 58: 10 (1875). - Micarea poliodes (Nyl.) Vezda in Vezda & V. Wirth in<br />
Folia geobot. phytotax., Praha 11: 99 (1976). Type: Finland, Tavastia australis, Evo, [on schistose rock<br />
with M. sylvicola],J. P. Norrlin (H-NYL 20683 - holotype!).
LICHEN GENUS MICAREA IN EUROPE 149<br />
Lecidea antrophila Larb. ex Leighton in Trans. Linn. Soc. London (Bot.) II, 1: 242 (1876). Type: Ireland,<br />
West Galway, Mwellan near Kylemore, in <strong>the</strong> interior <strong>of</strong> a cave, 1877, C. Larbalestier (BM ex K -<br />
lectotype!; isolectotypes: BM!, BM ex K!).<br />
Lecidea paucula Nyl. in Flora, Jena 59: 573 (1876). Type: Ireland, West Galway, 'Montagnes de Maam'<br />
[Maumturk Mountains], iii 1876, C. Lflr/?fl/e5ner(H-NYL 20090 -holotype!).<br />
Micarea umbrosa Vezda & V. Wirth in Folia geobot. phytotax., Praha 11: 93 (1976). Type: Germany,<br />
Baden, Siidschwarzwald, between Neuhausle and Altglashiitte near St. Margen, 830 m, 2 v 1969, V.<br />
Wirth (hb. Wirth 1609 - holotype!).<br />
Lecidea granvina Vainio in Havaas in Bergens Mus. Arb. 1909 (1): 31 (1909); nom. nudum (Art. 32). Spec.<br />
orig.: Norway, Hordaland, Hardanger, Granvin, 10 xi 1900,7. J. Havaas (BG!).<br />
Lecidea demarginata auct. p.p., non Nyl.; see 'Excluded Taxa'.<br />
Note: typification <strong>of</strong> Lecidea lutulata. Larbalestier's ga<strong>the</strong>rings in BM and H, that were collected from<br />
Rozel meadow in 1873 and subsequently labelled 'Lecidea lutulata', consist <strong>of</strong> M. bauschiana, M. lutulata<br />
and M. peliocarpa. The specimen H-NYL 10696 contains M. lutulata only and agrees with <strong>the</strong> original<br />
diagnosis, e.g. 'hypo<strong>the</strong>cium fusconigricans crassum', and is <strong>the</strong>refore selected as lectotype.<br />
Thallus effuse, thin (up to 40 /xm thick), ± smooth or finely rimose, usually becoming thicker<br />
(up to 600 jxm thick) and scurfy-granular (but never forming discrete areolae or goniocysts) pale<br />
,<br />
greenish grey or grey-green, sometimes straw-coloured in part, sometimes oxydated (on<br />
ferruginous rocks); in section ecorticate and without an amorphous hyaline covering layer.<br />
Phycobiont not micareoid; cells thin-walled, ± globose, c. 5-12 /xm diam, or ellipsoid and up to<br />
15x8/xm.<br />
Apo<strong>the</strong>cia numerous, convex-hemispherical and immarginate from <strong>the</strong> start, later becoming<br />
± globose or tuberculate, grey-brown, dark brown or blackish, always turning blackish when<br />
moistened, 0-2-0-4 mm diam, or up to 0-8 mm diam when tuberculate. Hymenium 30-40 ^im<br />
tall; upper part varying from hyaline or pale fuscous-brown through olivaceous to dark<br />
aeruginose, pigment <strong>of</strong>ten in patches corresponding to clustered apices <strong>of</strong> stout paraphyses,<br />
K- , HNO3+ red; lower part hyaline or dilutely coloured. Asci cylindrical-clavate, c. 30-40x7-<br />
10 /am. Spores ellipsoid, ovoid, or sometimes dacryoid, 6-8 (-9) x 2-3 (-3 -4) yam. Paraphyses<br />
ra<strong>the</strong>r scanty, <strong>of</strong> two types: p.p. evenly distributed, irregularly flexuose, simple, or forked or<br />
with short branches (especially in <strong>the</strong> upper part), sometimes anastomosing, 0-8-l(-l-5) ^tm<br />
wide, apices sometimes irregularly swollen to c. 2 ^tm wide; p.p. fasciculate, mostly simple,<br />
stout, c. 1-5-2 /xm wide, with swollen (to 3 /xm), coherent, pigmented apices. Hypo<strong>the</strong>cium<br />
120-360 /x,m tall, dark and opaque, fuscous- or reddish brown, K- or -I- reddish intensifying (but<br />
never with purple or greenish tinges), HNO3-; hyphae interwoven, but becoming vertically<br />
orientated towards <strong>the</strong> hymenium, c. 1-5-2 /xm wide but coated with dense brown pigment and<br />
appearing 3-4 /xm wide; ascogenous hyphae with swollen cells c. 2-4 /xm wide. Excipulum not<br />
seen even in sections <strong>of</strong> young apo<strong>the</strong>cia; <strong>the</strong> hymenium soon becomes reflexed so as to form an<br />
excipulum-like zone below <strong>the</strong> hypo<strong>the</strong>cium. Pycnidia <strong>of</strong>ten present but inconspicuous, immersed,<br />
blackish, c. 80-200 /xm diam, ostiole <strong>of</strong>ten gaping; with a single ± globose locule but<br />
walls <strong>of</strong>ten convoluted with up to 5 locules seen in a vertical section; outer walls reddish brown,<br />
K-; inner walls <strong>of</strong> secondary locules hyaline; conidiogenous cells irregularly cylindrical, <strong>of</strong>ten<br />
with 1-2 percurrent proliferations, 7-10x1-1-5 /xm, base sometimes swollen to 2-7 /xm wide;<br />
conidia (mesoconidia) cylindrical, sometimes faintly biguttulate but not constricted in <strong>the</strong><br />
middle, (3-8-)4-5(-5-5)x0-9-l-4/xm.<br />
Chemistry: All parts K-, C-, PD-; t.l.c: no substances detected.<br />
Observations: The apo<strong>the</strong>cia <strong>of</strong> M. lutulata have a similar internal pigmentation to those <strong>of</strong> M.<br />
botryoides {q. v. ) and M. muhrii {q. v. ), but <strong>the</strong> species is most <strong>of</strong>ten confused with M. sylvicola.<br />
The latter shares <strong>the</strong> same habitat as M. lutulata, but can be distinguished by its hypo<strong>the</strong>cium<br />
which always has a distinctly green, or rarely purplish, tinge in water mounts or KOH; in HNO3<br />
<strong>the</strong> pigment(s) turn purple-red. M. sylvicola fur<strong>the</strong>r differs in its larger spores, taller hymenium,<br />
slightly longer and broader conidia, and presence <strong>of</strong> greenish pigment in its pycnidial walls. M.<br />
myriocarpa occurs in similar habitats to M. lutulata and has a reddish brown, but never opaque<br />
(in thin section) hypo<strong>the</strong>cium. Fur<strong>the</strong>rmore, M. myriocarpa has smaller apo<strong>the</strong>cia, narrower,<br />
<strong>of</strong>ten 1-septate spores, and sessile pycnidia with much shorter conidia.
150<br />
BRIAN JOHN COPPINS<br />
Map 11 Micarea lithinella ® 1950 onwards + Micarea lutulata • 1950 onwards O Before 1950<br />
Habitat and distribution: M. lutulata is found on rocks and sometimes exposed roots in dry<br />
rocky underhangs, and is a characteristic species <strong>of</strong> <strong>the</strong> Micareetum sylvicolae. It is apparently<br />
widespread in nor<strong>the</strong>rn and western Britain; and although not yet reported from <strong>the</strong> south-west<br />
peninsula <strong>of</strong> England or <strong>the</strong> north-west <strong>of</strong> Scotland, it almost certainly occurs in those areas. It<br />
appears to be Uttle known outside <strong>of</strong> <strong>the</strong> <strong>British</strong> Isles, but I have seen material from Norway<br />
(Oppland and Hordaland), Sweden (Varmland), sou<strong>the</strong>rn Finland, Germany (Baden-<br />
Wiirttemberg), and Czechoslovakia (Bohemia: Krkonose).<br />
Exsiccata: Arnold Lich. Exs. 409B p.p. (WRSL). Havaas Lich. Exs. Norv. 571 (EG). Larbal. Lich.<br />
Herb. 223 (BM, BM ex K). Rasanen Lich. Fenn. 612 p. p. (LD-mixed with M. tuberculata)<br />
22. Micarea melaena (Nyl.) Hedl.<br />
'(Figs21A,46A;Mapl2)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 82, 96 (1892). - Lecidea melaena Nyl. in Bot. Notiser<br />
1853: 182 (1853). - Lecidea vernalis var. melaena (Nyl.) Nyl. in Mem. Soc. Imp. Sci. nat. Cherbourg. 3:<br />
182 (1855). - Bacidia melaena (Nyl.) Zahlbr. in Annls mycol. 7: 474 (1909). Type: Sweden, on lignum,<br />
E. M. Fries, Lich. Suec. Exs. 212B (UPS-lectotype!; isolectotypes: C!, H-NYLp.m. 4778 [fragment]!,<br />
M!).<br />
Lecidea milliaria var. turfosa Fr. , Nov. Sched. Crit. 8: 7 (1826), non Biatora turfosa Massal. Type: Sweden,<br />
[? Smaland], on peaty turf, E. M. Fries, Lich. Suec. Exs. 212A (UPS-lectotype!; isolectotypes: C!, M!).<br />
.
LICHEN GENUS MICAREA IN EUROPE 151<br />
Biatora stizenbergeri Hepp, Flecht. Eur. no. 504 (1860). Type: Switzerland, Rifferschwell, on dry plant<br />
debris amongst roots in peat moor, Hegetschweiler (BM - lectotype!; isotypes distributed in Hepp.<br />
Flecht. Eur. 504: BM!, E!, WRSL!).<br />
Lecidea ilyophora Stirton in Scott. Nat. 5: 220 (1879). Type: Scotland, Perthshire, Black Wood <strong>of</strong><br />
substances detected]).<br />
Rannoch, on lignum, ix 1879, J. Stirton BM - lectotype! [t.l.c. : no<br />
Lecidea melaena f. catillarioides Vainio in Medd. Soc. Fauna Fl. fenn. 10: 12 (1883). - Micarea melaena f.<br />
catillarioides (Vainio) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 83, 96 (1892). Type:<br />
Finland, Karelia borealis, Nurmes, Riihivaara, on bryophytes on rock, 1875, Vainio (TUR-VAINIO<br />
21478 -holotype!).<br />
Lecidea melaena f. endocyanea Vainio in Medd. Soc. Fauna Fl. fenn. 10: 12 (1883). - Micarea melaena f.<br />
endocyanea (Vainio) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill 18 (3): 83, 96 (1892). Type:<br />
USSR, Karelia keretina, Kivakka, on mosses on rocks, 1877, Vainio (TUR-VAINIO 21476 - hole-<br />
type!).<br />
Bilimbia melaena f. aeruginosa Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 254 (1922). Type: Finland,<br />
Tavastia australis, Hollola, on lignum, 1875, Vainio (TUR-VAINIO 21461 - lectotype!).<br />
Bilimbia melaena f. epiphaeotera Vainio in Acta Soc. Fauna Fl. Fenn. 53 (1): 255 (1922). Type: Finland,<br />
Karelia borealis, Nurmes, Louhivaara, 'Kallion syrjalla', on bryophytes and old Cladonia squamules,<br />
1875, Vainio (TUR-VAINIO 21472- lectotype!).<br />
?Bilimbia melaena var. alnicola Savicz in Izv. imp. S. Peterb. bot. Sada 14 (I, Suppl.): 53 (1914). Type:<br />
USSR (?LE); request for loan received no reply.<br />
Bilimbia milliaria y. [var.] saprophila Korber, Parerg. lich. : 171 (1860); nom. superft. (Art. 63).<br />
Thallus effuse, superficial, <strong>of</strong> small, scattered to coherent or clustered, granular areolae<br />
(20-)25-80(-100) /xm diam, sometimes forming an uneven crust up to c. 150 fxm thick. Areolae<br />
very variable in colour, pale buff or pale dull green to dark grey-green; in section without an<br />
amorphous covering layer, outermost hyphae <strong>of</strong>ten with dark green (K-, HNO3-I- red) walls.<br />
Thallus <strong>of</strong>ten scurfy granular and blackish (especially in damp situations) owing to its disruption<br />
by invading dematiaceous fungi and foreign algae; in dry situations (e.g. undersides <strong>of</strong> fallen<br />
trunks) it may have a whitish or pale yellowish , farinose appearance due to a thick covering <strong>of</strong> an<br />
epiphytic alga, each cell <strong>of</strong> which is coated by hyaline, crystalline incrustations. Phycobiont<br />
micareoid, cells c. 4-7 jxm diam.<br />
Apo<strong>the</strong>cia numerous, immarginate, convex-hemispherical to ± globose, sometimes tubercu-<br />
late, black, matt or slightly glossy, 0- 12-0-4 mm diam, or to 0-5 mm diam when tuberculate.<br />
Hymenium 30-40(-50) /xm tall, dull to bright aeruginose or grey-blue, K— or K-l- green<br />
intensifying; alternatively lower part (or rarely <strong>the</strong> entire hymenium) dilute purple-brown, K-l-<br />
greenish or K-l- purple intensifying; pigment(s) <strong>of</strong>ten more dense around <strong>the</strong> apices <strong>of</strong> <strong>the</strong><br />
paraphyses, thus appearing as a dark 'epi<strong>the</strong>cium' (c. 2-10 jxm tall) that contrasts with <strong>the</strong> more<br />
dilute colouration <strong>of</strong> <strong>the</strong> rest <strong>of</strong> <strong>the</strong> hymenium. Asci clavate, c. 30-40-12-15 jxm. Spores<br />
ovoid-oblong or oblong, with rounded apices, straight, (l-)3(-5)-septate, 12-21 x4-5(-5-5) /xm.<br />
Paraphyses numerous, branched and anastomosing, sometimes a few unbranched, thin, 0-8-1<br />
^im wide; apices not swollen, or sometimes slightly incrassate and up to 1-5 jxm wide, hyaline but<br />
<strong>of</strong>ten individually surrounded by a pigmented hood <strong>of</strong> dense gel (easily detached in mounts in K,<br />
by tapping <strong>the</strong> cover slip). Occasionally intermixed with ordinary paraphyses are a few which are<br />
simple , stout (c. 1 -5-2 jxm wide) and coated with pigment throughout <strong>the</strong>ir length so as to appear<br />
3-4 wide. Hypo<strong>the</strong>cium c. 80-160 ^im tall, dark and ± black in thick sections, pigmentation<br />
variable (see 'observations' below), usually purple-brown and K— or K-l- purple intensifying,<br />
or, outer (more rarely whole) part K-l- olivaceous with central part ('core') remaining ± purplish<br />
brown; all parts HNO3-I- purple-red; hyphae c. 0-7-1-5 jxm wide but <strong>the</strong>ir walls coated with<br />
dense pigment so that <strong>the</strong>y appear to be c. 3-4 /xm wide, interwoven or becoming vertically<br />
orientated towards <strong>the</strong> hymenium, sometimes a few extending into <strong>the</strong> hymenium as stout<br />
pigmented paraphyses; ascogenous hyphae similarly pigment, short-celled, c. 2-5 /am wide.<br />
Excipulum indistinct, sometimes evident in sections <strong>of</strong> young apo<strong>the</strong>cia, green or purplish<br />
(usually darker than hymenium, but paler than hypo<strong>the</strong>cium); hyphae radiating, branched and<br />
anastomosing, c. 0-7-1 jxm wide.<br />
Pycnidia rare, black, <strong>of</strong> two types: (a) c. 100-140 /xm diam, sessile; walls purple-brown or dull<br />
olive, K-l- green; conidia {macroconidia) curved or hamate, 0-7-septate, 18-33 x 1-1-5 /xm; (b)
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,
LICHEN GENUS MICAREA IN EUROPE 153<br />
-^y^<br />
Map 12 Micarea melaena # 1950 onwards O Before 1950<br />
C. floerkeana, C. subcervicornis, Coelocaulon aculeatum s. lat., Lecidea icmalea, Micarea<br />
leprosula, and Ochrolechia androgyna. M. melaena is <strong>of</strong>ten found on rocks, growing over peaty<br />
debris or decaying mats <strong>of</strong> bryophytes and Cladonia squamules, but occurrences where it is<br />
growing directly on rock are rare. To my knowledge, all such occurrences in Britain are confined<br />
to woodland situations in ra<strong>the</strong>r heavily polluted areas approximating to zones 3-5 on <strong>the</strong> scale<br />
<strong>of</strong> Hawksworth & Rose (1970); <strong>the</strong>se areas include Leicestershire, <strong>the</strong> West Yorkshire<br />
conurbation, and Fife.<br />
M. melaena is widely distributed in mainland Britain, but is particularly common in <strong>the</strong> upland<br />
areas <strong>of</strong> <strong>the</strong> north and west. There is a dearth <strong>of</strong> records from Ireland, where it is unlikely to be<br />
rare in suitable terrain. M. melaena is known from most parts <strong>of</strong> <strong>the</strong> European mainland and is<br />
particularly common in <strong>the</strong> boreal regions <strong>of</strong> Fennoscandia and upper Bavaria and adjoining<br />
alpine districts, but this apparent tendency may, at least partially, be a reflection <strong>of</strong> <strong>the</strong> avidity <strong>of</strong><br />
<strong>the</strong> collectors active in those areas, and <strong>the</strong> herbaria which I have studied! It has been found in<br />
<strong>the</strong> Azores on Cryptomeria and sulphur-rich soils near fumaroles, but it is so far unrecorded<br />
from <strong>the</strong> Canary Islands. From outside Europe I have seen several collections from eastern<br />
Canada and north-eastern U.S.A.<br />
Exsiccata: Anzi Lich. Exs. Ital. 259A (BM, M). Anzi Lich. Sondr. 170B (UPS). Arnold Lich. Exs.<br />
332A-C (M). Arnold Lich. Mon. 49 (M), 248 (C, M), 249 (C, MANCH), 407 (C, M, MANCH). Britz.<br />
Lich. Exs. 946 (M). Fellman Lich. Am. 159 (H). Flotow Lich. Exs. 129C (UPS). Fries Lich. Suec. 212A, B<br />
(C, H-NYL p.m. 4778, M, UPS). Hepp Fl. Eur. 504 (BM, E, WRSL). Johnson Lich. Herb. 376 (BM).<br />
Krypt. Exs. Vindob. 362 (BM, M). Leighton Lich. Brit. 120 (BM). Malme Lich. Suec. 27 (C, M, S). Moug.
154 BRIAN JOHN COPPINS<br />
& Nestl. Stirp. Crypt. 1329 p.p. (E). Mudd Lich. Brit. 159 (M, MANCH). Norrlin & Nyl. Herb. Lich.<br />
Fenn. 180 (BM, C, M). Oliv. Lich. Orne 237 (M). Rasanen Lich. Fenn. 963 (M). Rasanen Lichenoth.<br />
Fenn. 344 (BM). Roum. Lich. Gall. 231 (M). Schaerer Lich. Helv. 196 p.p. (E). Vezda Lich. Set. 14 (BM,<br />
M). Zwackh Lich. Exs. 657, 675 (M).<br />
23. Micarea melaenida (Nyl.) Coppins, comb. nov.<br />
(Fig. 22)<br />
Lecidea melaenida Nyl. in Flora, Jena 48: 146 (1865). - Catillaria melaenida (Nyl.) Arnold in Flora, Jena<br />
53: 475 (1870). Type: France, [? Eure], Perrieres, Lenormand (H-NYL 18843 -holotype!).<br />
Lecidea relicta Nyl. in Flora, Jena 48: 354 (1865) [nee Stirton (1876)]. Type: France, Calvados, Falaise,<br />
L.-A. De Brebisson (H-NYL 21099 - holotype!).<br />
Catillaria schumannii {'SchumannV] Stein in Cohn, Krypt. -Fl. Schles. 2 (2): 232 (1879). - Lecidea<br />
schumanni Korber, Syst. Lich. Germ.: 255 (1855); nom. inval. (Art. 34). Type: East Germany,<br />
Reichenbach, near Ernsdorf, Schumann (WRSL - lectotype!; WRSL - isolectotype!).<br />
Catillaria zsakii Szat. in Magy. hot. Lap. 24: 108 (1926). - Toninia zsakii (Szat.) Lettau in Feddes Reprium<br />
Spec. nov. veg. SI: 9 (1955). Type: holotype not seen (? in BP); topotypes: Hungary, 'In argillosis<br />
natronatis prope oppidum Karcag, Comit. Jasz-Nagykun-Szolnok. Allit. ca. 80 m. s.m. Mens. Szept.<br />
1926', Z. Zsdk cfe G. Timko, Fl. Hung. Exs. 714 (E! [t.l.c: no substances], BM!, BM ex K!), and, from<br />
same locality but at '100 m. s.m.' and no date, Krypt. Exs. Vindob. 3154 (BM!, BM ex K!).<br />
Catillaria schumanni var. meridionalis Roux & Vezda in Vezda, Sched. Lich. Sel. Exs. 537 (1967). Type:<br />
France, Gard, Pujaut near Avignon, alt. 100 m, 'supra solum argillaceo-sabulosum, ad marginem<br />
Querceti ilicis', 6 xi 1966, C. Roux, Vezda Lich. Sel. Exs. 537 (BM - isotype!).<br />
Thallus effuse, terricolous (on argillaceous soils), forming a thin ± smooth, dull greenish<br />
white crust, c. 40-100 /xm thick, or forming confluent, convex areolae 0- 1-0-6 mm diam; crust<br />
sometimes becoming cracked so as to appear ± 'squamulose'. Thallus in section without a<br />
distinct hyaline, amorphous, covering layer, but with a hyaline 'cortex' 12-20 fxm tall, composed<br />
<strong>of</strong> compacted, evacuated hyphae (cytoplasm presumed lost owing to lack <strong>of</strong> staining by LCB or<br />
ammoniacal erythrosin); thallus hyphae l-5-2-5(-3) fxm wide; numerous algae-free hyphae<br />
penetrating <strong>the</strong> soil substratum to a depth <strong>of</strong> c. 1 mm (? or more). Phycobiont micareoid, cells<br />
4-7 /Lim diam.<br />
Apo<strong>the</strong>cia numerous, scattered or 2-4-confluent, immarginate, convex to hemispherical,<br />
black, matt, 0-2-0-6 mm diam; old, contorted apo<strong>the</strong>cia sometimes reaching 1-2 mm diam;<br />
occasionally a fine weft <strong>of</strong> radiating, white (hyaline) hyphae is seen around <strong>the</strong> edge <strong>of</strong> an<br />
apo<strong>the</strong>cium. Hymenium 35-40 ^tm tall; uppermost c. 1-5 /am (epi<strong>the</strong>cium) with dense,<br />
purple-red; remaining (lower) parts dilute<br />
dull purple, K— , HNO3+ red. Asci clavate, 35-38x9-12 /xm. Spores ellipsoid, oblong-ellipsoid<br />
or ovoid-oblong, straightor slightly curved, 0-1-septate, (7-)9-5-15x3^(-4-5) /am. Paraphyses<br />
numerous, mostly simple below, but <strong>of</strong>ten branched above, sometimes anastomosing, 1-1-7 ixm<br />
wide; apices (upper 5-15 /xm) <strong>of</strong>ten purple-brown pigmented and to 2-5 /xm wide. Hypo<strong>the</strong>cium<br />
amorphous, purple-brown pigment, K— , HNO3+<br />
60-120(-200) /Ltm tall, mottled dark purplish brown, K— , or K+ purple intensifying in part,<br />
HNO3-I- purple-red; hyphae interwoven but ± vertically orientated in uppermost part, c. 1-1-8<br />
/am wide, each with a densely pigmented gel sheath <strong>the</strong>reby giving a total diameter <strong>of</strong> c. 3 /xm;<br />
ascogenous hyphae c. 2-5-5 /am wide. Excipulum soon reflexed, but distinct in sections <strong>of</strong> young<br />
apo<strong>the</strong>cia, purple-brown (usually darker than hymenium); hyphae radiating, branched and<br />
anastomosing, c. 1-3-2 /xm wide, embedded in densely pigmented gel-matrix.<br />
Pycnidia <strong>of</strong>ten present, immersed, black, c. 50-100 /xm diam; walls purple-brown, K— or Kf-Ipurple<br />
intensifying; conidia (? microconidia) , ± cylindrical, 4-8-6(-7)xl /xm.<br />
Chemistry: Thallus K— , C— , KC—<br />
, PD-; t.l.c: no substances.<br />
Observations: Micarea melaenida is characterized by its whitish, terricolous thallus, immar-<br />
ginate, black apo<strong>the</strong>cia (with purplish pigmentation in section), numerous and ra<strong>the</strong>r stout<br />
paraphyses, and 0-1-septate spores. It is apparently related to M. assimilata {q. v. ) and its allies.<br />
When compared with M. melaenida, forms <strong>of</strong> M. melaena with immature spores can be<br />
distinguished by <strong>the</strong>ir finely granular areolae, narrow and less numerous paraphyses, and usual
LICHEN GENUS MICAREA IN EUROPE 155<br />
presence <strong>of</strong> green pigments in apo<strong>the</strong>cial sections (in water or K mounts). M. turfosa is similar in<br />
some ways to M. melaenida, but has a dark coloured thallus, an olivaceous hymenium, reddish<br />
brown (never purplish) hypo<strong>the</strong>cium, and larger spores which are <strong>of</strong>ten 1-3-septate.<br />
Habitat and distribution: M. melaenida grows on, and in direct contact with, fine-grained<br />
sandy or argillaceous, mineral soils, and is not known to occur on decaying bryophytes or plant<br />
debris, etc. Unfortunately I have never seen M. melaenida in <strong>the</strong> field, and <strong>the</strong>re is little<br />
ecological information to be gleaned from <strong>the</strong> literature or herbarium labels. Fur<strong>the</strong>rmore, <strong>the</strong><br />
specimens examined have included few associated species, and only Cladonia sp. (squamules)<br />
and Catapyrenium lachneum have been noted. In its Hungarian locality it was said to have<br />
occurred on soil rich in precipitated salts ('argillosis natronalis') although no effervescence was<br />
apparent when I tested <strong>the</strong> soil with 50% HNO3. It is probable that M. melaenida is restricted to<br />
niches where <strong>the</strong> salts have been permanently leached out. A sample <strong>of</strong> soil from Fl. Hung. Exs.<br />
714 was sent for analysis to <strong>the</strong> Macaulay Institute for Soil Research (Aberdeen) and Dr B. W.<br />
Bache <strong>of</strong> <strong>the</strong> Institute reports 'It bears no relationship at all to a sodic soil, and is in fact fairly<br />
acid, with pH 5-2. The conductivity is fairly low (0-8 mS in 1:2 extract), so it does not contain any<br />
great concentration <strong>of</strong> soluble salts, nor does it contain any carbonate.'<br />
The data on <strong>the</strong> herbarium labels, although limited, do indicate that it occurs at low altitudes,<br />
for example <strong>the</strong> collection from Loire Atlantique was made from soil on coastal cliffs; that from<br />
Gard was made at 100 m; and those from Hungary were made at 80-100 m.<br />
M. melaenida is a rare species with a ra<strong>the</strong>r sou<strong>the</strong>rn distribution, and I have seen material<br />
from several localities in France (e.g. Loire Atlantique, Vienne, and Gard), and single localities<br />
in East Germany (Dresden) and Hungary. It is not known from Scandinavia, and its status as a<br />
<strong>British</strong> plant is doubtful. Its first <strong>British</strong> report seems to be its appearance in <strong>the</strong> checklist <strong>of</strong><br />
James (1965fl), but I have been unable to ascertain <strong>the</strong> reason for its inclusion. However, its<br />
presence in north-west France suggests that it could occur in Britain, especially along <strong>the</strong> south<br />
coast <strong>of</strong> England or in <strong>the</strong> Channel Islands.<br />
Material from South Africa and distributed as Almborn Lich. Africani no. 84 is not M.<br />
melaenida but a mixture <strong>of</strong> two, undescribed species, both <strong>of</strong> which are probably referable to<br />
Micarea.<br />
Exsiccata: Fl. Hung. Exs. 714 (BM, BM ex K, E). Krypt. Exs. Vindob. 3154 (BM, BM ex K). Vezda<br />
Lich. Sel. 537 (BM).<br />
24. Micarea melaenizaHedl.<br />
(Figs21B,38B)<br />
in Bih. K. svenska VetenskAkad. Hand. Ill, 18 (3): 83, 96 (1892). - Lecidea melaeniza ["melaniza']<br />
(Hedl.) Zahlbr., Cat. lich. univ. 3: 798 (1925). Type: Sweden, Halsingland, lignum <strong>of</strong> old pine trunk, viii<br />
1891,7. T. Hedlund iS-ho\otype\).<br />
Thallus effuse, lignicolous (sometimes extending on to bryophyte or lichen thalU), endoxyhc,<br />
or epixylic with whitish convex areolae; areolae c. 100-400 fxm diam and up to 45 /am thick,<br />
without an amorphous covering layer. Phycobiont micareoid, cells 4-7 /xm diam.<br />
Apo<strong>the</strong>cia numerous, few, or absent, immarginate, at first convex-hemispherical or subglobose,<br />
<strong>of</strong>ten becoming tuberculate, black, matt, 0-2-0-3 mm diam, or to 0-5 mm diam when<br />
tuberculate. Hymenium 28-42 /xm tall, upper and lower parts aeruginose-green, middle part ±<br />
hyaline with aeruginose vertical streaks; K- or dulling, HNO3-I- red. A^dclavate, 26-35 x 10-12<br />
yarn. Spores ovoid or dacryoid, simple, 5-9x2- 5-3 -8 yam. Paraphyses ra<strong>the</strong>r numerous, <strong>of</strong> two<br />
types; p.p. evenly distributed, flexuose, branched, thin, c. 0-7-1 ^tm wide, hyahne, but apices<br />
sometimes pigmented and <strong>the</strong>n to 2-5 /am wide; p.p. scattered or in small fascicles, stout, c. 2 /am<br />
wide, each coated ± throughout <strong>the</strong>ir length by greenish pigment. Hypo<strong>the</strong>cium c. 120-200 /am<br />
tall, dark brown, sometimes with a faint reddish (never purplish) tinge, K- or dulling, HNO3 -<br />
or red tinge slightly intensifying; hyphae coated with dark brown pigment and c. 2-3 /am wide,<br />
interwoven but becoming vertically orientated towards <strong>the</strong> hymenium and sometimes con-
156 BRIAN JOHN COPPINS<br />
tinuing into it as stout pigmented paraphyses; ascogenous hyphae similarly pigmented, but with<br />
short swollen cells to 5 /xm wide. Excipulum not evident in available material.<br />
Pycnidia numerous, black, sessile and c. 40-60 /xm diam, or more usually borne on black<br />
stalks (pycnidiophores) and 80-300x40-70 ^im (including stalk); stalks simple, or sometimes<br />
branched and bearing up to four pycnidia; upper part (pycnidial wall) dark olive but lower part<br />
(stalk tissue) reddish brown, both parts K— , Conidia {mesoconidia) short cylindrical, 2-3-<br />
3-6xl-l-3/>im<br />
Chemistry: Thallus PD— ; sections <strong>of</strong> apo<strong>the</strong>cia and thallus C-; material insufficient for<br />
analysis by t.l.c.<br />
Observations: With its black, subglobose to tuberculate apo<strong>the</strong>cia, black, stalked pycnidia,<br />
inconspicuous thallus, and occurrence on lignum, M. melaeniza is indistinguishable in external<br />
appearance from M. misella and M. nigella. Microscopically, M. misella can be distinguished by<br />
<strong>the</strong> olivaceous (K+ violet) pigment in its hymenium and pycnidia, and its ± hyaline hypo<strong>the</strong>cium;<br />
and M. nigra can be distinguished by <strong>the</strong> purple-brown (K+ green, HNO3+ purple-red)<br />
pigment in its hymenium, hypo<strong>the</strong>cium and pycnidia. With respect to apo<strong>the</strong>cial and pycnidial<br />
pigmentation and structure, M. melaeniza is almost identical to M. botryoides. However, M.<br />
botryoides is rarely lignicolous, and has larger, <strong>of</strong>ten septate spores, and longer mesoconidia. M.<br />
muhrii has similar apo<strong>the</strong>cial pigments to those <strong>of</strong> M. melaeniza, but is readily distinguished by<br />
its larger, adnate apo<strong>the</strong>cia (which <strong>of</strong>ten have a pale marginal rim), larger spores, well<br />
developed excipulum, and lack <strong>of</strong> stalked pycnidia.<br />
Habitat and distribution: M. melaeniza is a rare species, known only from three collections<br />
made in Sweden (Angermanland, Halsingland, and Smaland) where it occurred on <strong>the</strong> lignum<br />
<strong>of</strong> conifer trunks. Associated species identified on <strong>the</strong> examined collections include Calicium<br />
salicinum, Cetraria pinastri, Chaeno<strong>the</strong>copsis lignicola, Cladonia spp. (squamules), Dimerella<br />
diluta, Hypogymnia physodes, Lecanora symmicta agg., Micarea anterior, M. misella, M.<br />
prasina, Usnea sp., and Lophozia sp.<br />
25. Micarea melanobola (Nyl.) Coppins, comb. nov.<br />
(Figs 21c, 46B-C)<br />
Lecidea melanobola Nyl. in Flora, Jena 50: 371 (1867). - Catillaria melanobola (Nyl.) Vainio in Acta Soc.<br />
Fauna Fl. fenn. 57 (2): 465 (1934). - Lecidea erysiboides *L. melanobola (Nyl.) Nyl. in Hue in Revue Bot.<br />
Courrensan 5: 103 (1886). - Catillaria |3. byssacea f. melanobola (Nyl.) Blomb. & Forss., Enum. PI.<br />
Scand. : 91 (1880). - Micarea prasina f . melanobola Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18<br />
(3): 87 (1892). Type: Finland, Tavastia australis, Kuhmois [Kuhomoinen], on bark <strong>of</strong> Picea abies ,<br />
/. P. Norrlin (H-NYL 21614 -lectotype!; isolectotypes: H-NYLp.m. 4504!, H!).<br />
Thallus effuse, thin and indistinct, <strong>of</strong> scattered to ± coherent, minute olivaceous granules<br />
(goniocysts), c. 12-25 jxm diam, which appear ± gelatinous when moist. Outermost hyphae <strong>of</strong><br />
goniocysts with greenish, K-l- violet walls. Phycobiont micareoid, cells 4-7 (xm diam.<br />
Apo<strong>the</strong>cia numerous, convex-hemispherical, immarginate, dark grey to black, 0- 1-0-24 mm<br />
diam. Hymenium 30-35 /xm tall, hyaline, but with a clearly delimited, dark green epi<strong>the</strong>cium,<br />
K+ deep violet, C+ violet, HNOs-l- red; pigment closely bound to (? or also in) <strong>the</strong> apical walls<br />
<strong>of</strong> <strong>the</strong> paraphyses, and also present in <strong>the</strong> gel-matrix and apical walls <strong>of</strong> <strong>the</strong> asci. Asci clavate,<br />
30-35x9-11 fjLm, upper wall(s) sometimes tinged with greenish, K+ violet pigment. Spores<br />
ellipsoid, ovoid, or oblong-ovoid, with obtuse apices, 0-1-septate, 7-9-7x2-5-3-3 /xm. Paraphyses<br />
scanty, branched and anastomosing, thin, 0-5-1 /xm wide; apices thickened with greenish<br />
(K-l- violet) pigment and up to 1-7 /xm wide, <strong>of</strong>ten overtopping <strong>the</strong> tops <strong>of</strong> <strong>the</strong> asci. Hypo<strong>the</strong>cium<br />
25-70 /xm tall, hyaline or dilute dull yellowish. Excipulum indistinct, sometimes evident as a<br />
non-amyloid, narrow (c. 10-15 /xm), reflexed zone <strong>of</strong> radiating, branched and anastomosing<br />
hyphae, 0-5-1 /xm wide.<br />
Pycnidia numerous but small and inconspicuous, sessile, black, with dark greenish (K-l-<br />
violet) walls; <strong>of</strong> two types: (a) 40-50 /xm diam; conidia (mesoconidia) cylindrical or ovoid-<br />
1866,<br />
I
LICHEN GENUS MICAREA IN EUROPE 157<br />
Table 6 Diagnostic features for <strong>the</strong> separation <strong>of</strong> Micarea melanobola, M. misella, and M. prasina.
158 BRIAN JOHN COPPINS<br />
26. Micarea misella (Nyl.) Hedl.<br />
(Fig. 23A; Map 13)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 88 (1892). - Lecidea anomala f. misella Nyl.,<br />
202 (1861) .- Lecidea misella (Nyl.) Nyl. in Not. Sdllsk. Fauna Fl.fenn. Fork. 8: 177 (1866).<br />
Lich. Scand. :<br />
Type: Finland, Nylandia, Helsingfors [Helsinki], 1858, W. Nylander (H-\ectotypel).<br />
Lecidea resinae* [subsp.] L. globularis Nyl. , Lich. Scand. : 213 (1861). - Lecidea globularis (Nyl.) Lamy in<br />
Bull. Soc. bot. Fr. 25: 435 (1878). - Micarea globularis (Nyl.) Hedl. in Bih. K. svenska VetenskAkad.<br />
Handl. Ill, 18 (3): 78, 88 (1892). Type: Sweden, 'Arthonia turgida var. y globifera Svecia 112°' (H-ACH<br />
52-holotype!).<br />
Lecidea melanochroza Leighton ex Crombie in 7. Bot., Lond. 9: 178 (1871). Type: Scotland, Perthshire,<br />
near Loch Tummel, /. M. Crombie (BM - lectotype!; isolectotypes: BM ex K!, and distributed in<br />
Crombie Lich. Brit. Exs. 174 (BM!) [note: this example includes M. prasina and those in E and M<br />
contain M. prasina only]).<br />
Lecidea misella f. brasiliana Vainio in Acta Soc. Fauna Fl. fenn 7 (2): 57 (1890). Type: Brazil, Minas<br />
Geraes, Carassa, 1400 m, on lignum, Vainio, Lich. Brasil. Exs. 1420 (BM ex K - lectotype!; M -<br />
isolectotype!).<br />
Bilimbia melaena f. decrustata Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 255 (1922). Type: Finland,<br />
Tavastia australis, HoUola, 1871, Vainio (TUR-VAINIO 21480- lectotype!).<br />
Lecidea asserculorum sensu Th. Fr., Lich. Scand. 2: 473 (1874), non Ach (1810). See note by Hedlund<br />
(1892: 89).<br />
Thallus effuse, <strong>of</strong>ten wide-spreading, usually lignicolous and endoxylic, sometimes develop-<br />
ing on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> substratum as convex to irregularly subglobose, greenish white to<br />
greenish grey, ± contiguous granular-areolae, c. 60-120(-150) /w-m diam. Areolae in section,<br />
without a well defined cortex or hyaline amorphous covering layer, but outer 5-12 /xm is <strong>of</strong>ten<br />
tinged with olivaceous-brown pigment (K+ violet); pigment confined to gel-matrix and not<br />
adhering to hyphal walls. Phycobiont micareoid, cells 4—7 /xm diam.<br />
Apo<strong>the</strong>cia numerous, few or absent, scattered to crowded and confluent, immarginate,<br />
convex-hemispherical to subglobose, sometimes shortly stipitate, sometimes tuberculate, black,<br />
rarely pallid or grey-brown (shade forms), matt. c. 0-1-0-3 mm diam. Hymenium 25-36 /am tall;<br />
upper part dull greenish brown or olivaceous, K-l- violet, with pigment confined to <strong>the</strong><br />
gel-matrix; remaining (lower) part hyaline or dilute sordid greenish. Asci clavate, 25-35x7-10<br />
^tm. Spores ellipsoid, ovoid, or oblong-ovoid, not curved, sometimes with 1-3 guttules, simple<br />
but a few (very rarely many) sometimes 1-septate, (6-5-)7-9-5x2-0-3(-3-7) jxm. Paraphyses<br />
scanty, sparingly branched 0-5-0-8 /xm wide; apical parts <strong>of</strong>ten more richly branched, sometimes<br />
wider (to 1-5 /xm), but without closely adhering pigment. Hypo<strong>the</strong>cium c. 45-70 /xm tall,<br />
hyaline, or dilute olivaceous (rarely dark olivaceous: 'f. brasiliana') and <strong>the</strong>n K-f- violet; hyphae<br />
interwoven but becoming vertically orientated towards <strong>the</strong> hymenium, c. 0-7-1 /xm wide;<br />
ascogenous hyphae with swollen cells, c. 2-4 /xm wide. Excipulum poorly developed but <strong>of</strong>ten<br />
evident as a dark brownish (K+ violet) zone bordering <strong>the</strong> reflexed edges <strong>of</strong> <strong>the</strong> hymenium;<br />
hyphae radiating, branched and anastomosing, very narrow, c. 0-5-0-8 /xm wide.<br />
Pycnidia usually present black, walls green-brown, K-l- violet; <strong>of</strong> two types: (a) sometimes<br />
sessile, more usually borne single in <strong>the</strong> top <strong>of</strong> a black, unbranched stalk (pycnidiophore),<br />
70-320 /xm tall (including stalk) and 50-100 /xm wide; stalk part composed <strong>of</strong> interwoven<br />
hyphae, c. 1-1 -5 /xm wide, embedded in a gel-matrix containing a dull brown or olivaceous, K-lviolet<br />
pigment; conidia (mesoconidia) shortly cylindrical, sometimes faintly biguttulate, 3-5-<br />
5xl-l-5(-l-7) /xm; (b) ± immersed in <strong>the</strong> substratum (lignum) or areolae, c. 40 /xm diam; wall<br />
with green-brown, K-l- violet pigment in upper part, but becoming ± hyaline below; conidia<br />
(microconidia) narrowly cylindrical, 3-8-6x0-6--0-8 /xm. Pycnidia with stalks (a) are mostly<br />
confined to forms with an endoxylic thallus.<br />
Chemistry: Sections <strong>of</strong> thallus (when superficial) K- , C-l- red, PD- ; sections <strong>of</strong> apo<strong>the</strong>cia C-fviolet<br />
(olivaceous pigment), but only rarely C-l- red; t.l.c: gyrophoric acid detected in varying<br />
amounts, sometimes absent.<br />
Observations: Typical forms <strong>of</strong> M. misella with an endoxylic thallus, numerous apo<strong>the</strong>cia, and<br />
stalked pycnidia are easy to recognize. However, when it has a well developed superficial thallus
LICHEN GENUS MICAREA IN EUROPE<br />
Table 7 Diagnostic features for <strong>the</strong> separation <strong>of</strong> Micarea denigrata and M. misella.<br />
Apo<strong>the</strong>cia size (mm)<br />
Thallus type<br />
Spore septation<br />
Spore length (/u.m)<br />
Paraphyses<br />
Hymenium<br />
Macroconidia (/u,m)<br />
Mesoconidia (/Am)<br />
Microconidia (/u,m)<br />
Pycnidia (meso-)<br />
Chemistry (t.l.c.)<br />
0-1-0-3<br />
Usually endoxylic; sometimes<br />
with areolae, 60-120 /x,m diam<br />
0(-l)<br />
6-5-9-5<br />
sparse; 0-5-0-8 )u,m wide<br />
C+ violet; very rarely also<br />
C+ orange-red<br />
unknown<br />
3-5-5 X 1-1-5<br />
3-8-5x0-6-0-8<br />
sessile to stalked<br />
nil; but gyrophoric acid<br />
sometimes present in trace<br />
amounts, rarely in large amounts<br />
misella denigrata<br />
0-1-0-5<br />
usually with areolae, 60-200 /nm<br />
diam; rarely endoxylic<br />
(0-)l<br />
(7-)9-16(-18)<br />
numerous; 1-1-7 jxm wide<br />
C+ violet; usually also<br />
C-l- orange-red<br />
12-24x1; curved<br />
2-8-4-5(-5)x 1-2-1-8<br />
(4-5-)5-7-5x 0-7-0-8<br />
immersed to emergent, never<br />
stalked<br />
gyrophoric acid usually present<br />
in large amounts; very rarely<br />
not detectable<br />
and no stalked pycnidia it can be difficult to separate from M. denigrata, especially if in <strong>the</strong> latter<br />
no conidial (or only <strong>the</strong> microconidial) state can be found. In such instances M. misella can<br />
usually be distinguished by its small apo<strong>the</strong>cia, mainly simple, uncurved ascospores and sparse,<br />
thin paraphyses. The same criteria can separate forms <strong>of</strong> both species with an endoxylic thallus<br />
and lacking diagnostic macroconidial or mesoconidial states. A C+ orange-red reaction can<br />
usually be obtained in sections <strong>of</strong> <strong>the</strong> apo<strong>the</strong>cia <strong>of</strong> M. denigrata, although <strong>the</strong> reaction is<br />
sometimes very weak or unobtainable in diminutive, lignicolous, epixylic forms; in M. misella I<br />
have obtained this reaction in only one or two extremely well-developed specimens. Table 7<br />
outlines <strong>the</strong> main diagnostic features <strong>of</strong> <strong>the</strong>se two species. Apart from M. misella, black, stalked<br />
pycnidia are also a characteristic feature <strong>of</strong> M. botryoides, M. melaeniza, and M. nigella;<br />
however, <strong>the</strong> pycnidial wall tissue in <strong>the</strong>se species is brownish or greenish in KOH, and never<br />
violaceous as in M. misella.<br />
For differences from M. melanobola and M. prasina see Table 6.<br />
Habitat and distribution: M. misella is almost exclusively lignicolous (especially on conifer<br />
lignum). It has not been found on bark, but has been collected on Picea cones, old basidiomes <strong>of</strong><br />
polypores (Daedaleopsis confragosa and Gloeophyllum sepiarum), and on moribund Polytrichum<br />
spp. by woodland tracks. Most <strong>British</strong> collections <strong>of</strong> M. misella have been made from<br />
<strong>the</strong> sides and undersides <strong>of</strong> fallen decorticated trunks <strong>of</strong> Pinus, and associated species included<br />
Lecidea icmalea, Micarea prasina, Xylographa abietina, X. vitiligo, Cladonia spp. (scattered<br />
squamules), and Hypogymnia physodes. Reports on exposed roots and stones (e.g. Watson,<br />
1930: 53) refer to o<strong>the</strong>r species, especially M. bauschiana. In Britain M. misella appears to be<br />
ra<strong>the</strong>r rare and restricted to Scotland where it is mainly found in <strong>the</strong> native pine-woods. It has<br />
been reported (as 'Lecidea asserculorum') from England and Wales, but all <strong>the</strong> pertinent<br />
material seen belongs to diminutive forms <strong>of</strong> M. denigrata and M. prasina, or quite different<br />
species such as M. melaena and Cliostomum grifftthii.<br />
M. misella is widely distributed in Europe, especially in areas with naturally occurring<br />
coniferous forests. It is found as far north as Lycksele Lappmark and Norbotten in Sweden, and<br />
an outlying sou<strong>the</strong>rly locality is on <strong>the</strong> Mediterranean island <strong>of</strong> Corsica where it was collected on<br />
<strong>the</strong> rotting lignum <strong>of</strong> Eucalyptus. From outside Europe I have seen material <strong>of</strong> M. misella from<br />
Canada (Ontario and Newfoundland), and from <strong>the</strong> highlands <strong>of</strong> Minas Geraes (at 1400 m) in<br />
Brazil.<br />
Exsiccata: Arnold Lich. Exs. 626, 627 (BM ex K, M). Arnold Lich. Mon. Ill, 241, 307 (BM ex K, M,<br />
MANCH). Britz. Lich. Exs. 208 (M). Claudel «& Harm. Lich. Gall. 445 (BM). Crombie Lich. Brit. 11<br />
A<br />
159
160 BRIAN JOHN COPPINS<br />
Map 13 Micarea misella # 1950 onwards O Before 1950<br />
p.p. (BM). Elenkin Lich. Ross. 189 (UPS). Krypt. Exs. Vindob. 1532 (BM, BM ex K, M), 4214 (BM ex K,<br />
M). Malme Lich. Suec. 365 (S, WIS). Norrlin & Nyl. Herb. Lich. Fenn. 744 (H). Reliq. Suza. 42 (BM,<br />
GZU, WIS). Vainio Lich. Bras. 1420 (BM ex K, M), 1451 (BM). Zwackh Lich. Exs. 1085 (M).<br />
27. Micarea muhrii Coppins, sp. nov.<br />
(Fig. 23B)<br />
Thallus effusus, endoxylicus et labem albidam efficiens, vel epixylicus aut epilithicus; thallus superficialis<br />
cum areolis convexis, albidus, c. 0-06-0.2 mm diam, interdum coalescentibus in crustam crassiusculam<br />
demum rimosam. Algae cellulis 4-7 yam diam. Apo<strong>the</strong>cia adnata, convexa vel hemisphaerica autem baud<br />
globosa, griseo-atra, aut pallida vel rufo-brunneola vel fumosa ubi umbrose, 0- 15-0-46 mm diam,<br />
immarginata sed vulgo zona marginali, complana, ± incolorata. Hymenium 40-45 jxxn altum, parte summa<br />
obscure olivacea vel griseo-viridi et K— , vel raro hyalina. Ascosporae ellipsoideae, ovoideo-ellipsoideae<br />
vel oblongo-ellipsoideae, simplices, 9-12x4-5 /u,m. Paraphyses aliquantum paucae, ramosae et interdum<br />
anastomosantes, 1-1-5 /u,m latae, apices versus vulgo incrasstae ad 3-5 /xm latae et vulgo pigmentiferae.<br />
hyalinum, paulum evolutum mox reflexum.<br />
Hypo<strong>the</strong>cium obscure ruf<strong>of</strong>uscum, K— , HNO3—<br />
. Excipulum<br />
Pycnidia pauca, immersa. Conidia cylindrica, 4-6x0-8-1 /x.m. Thallus et apo<strong>the</strong>cia K-, C-, PD — ; sine<br />
materia chemica.<br />
Typus: Suecia, Wermelandia, par. Lungsund, Pungbacken, alt. c. 170 m, in loco aperto ad truncum<br />
decorticatum supra rivum, 15 vii 1980, leg. L.-E. Muhr 2851 (E-holotypus). [t.l.c: no substances.]
LICHEN GENUS MICAREA IN EUROPE 161<br />
Thallus effuse, endoxylic and visible as a white stain, or epixylic, or rarely epilithic; superficial<br />
thallus formed <strong>of</strong> whitish, greenish-white or grey-white, convex areolae, c. 0-06-0-2 mm diam,<br />
which are especially well developed around <strong>the</strong> apo<strong>the</strong>cia; areolae sometimes coalescing to form<br />
a thickish crust up to 0-2 mm thick, which may eventually become rimose. Areolae in section<br />
without a hyaline amorphous covering layer. Phycobiont micareoid, cells 4-7 /u,m diam.<br />
Apo<strong>the</strong>cia numerous, occasionally coalescing, adnate, convex, soon becoming hemispherical,<br />
but never globose or tuberculate, grey-black, or pallid, reddish brown or brown-grey in shade<br />
forms, 0- 15-0-46 mm diam; disc matt or slightly glossy; immarginate but <strong>the</strong> adnate rim is <strong>of</strong>ten<br />
seen as a ± colourless or watery grey zone, c. 20-40 /xm wide. Hymenium 40-45 /xm, ± hyaUne<br />
with upper part dark olivaceous or grey-green, K— , HNOs-f- purple-red, or ± hyaline<br />
throughout in shade forms. Asci cylindrical-clavate, 38-40x10-12 /xm. Spores simple, <strong>of</strong>ten<br />
biguttulate, ellipsoid, ovoid-ellipsoid or oblong-ellipsoid, 9-12x4-5 /xm. Paraphyses ra<strong>the</strong>r<br />
scanty, branched (especially in <strong>the</strong>ir upper parts), sometimes anastomosing, 1-1-5 /zm wide,<br />
<strong>of</strong>ten sUghtly incrassate at apices to 2 /^m wide; or upper 9-15 /xm <strong>of</strong> paraphyses thickened by<br />
dense greenish pigment and <strong>the</strong>n to 3-5 /xm wide. Hypo<strong>the</strong>cium c. 70-120 ^im tall, dark reddish<br />
brown, K— , HNO3—<br />
; hyphae c. 1-2 /xm wide embedded in <strong>the</strong> densely pigmented matrix,<br />
interwoven but becoming vertically orientated towards <strong>the</strong> hymenium; ascogenous hyphae with<br />
short, swollen cells to 5 (xm wide. Excipulum sometimes evident in sections <strong>of</strong> young apo<strong>the</strong>cia<br />
as a non-amyloid, hyaline zone, but soon reflexed and obscured in older apo<strong>the</strong>cia; composed <strong>of</strong><br />
radiating, branched and anastomosing hyphae l-l-5(-2) ^im wide.<br />
Pycnidia rare, sunken within <strong>the</strong> areolae, c. 40-50 fxm diam; upper part <strong>of</strong> wall around <strong>the</strong><br />
ostiole olivaceous, lower part reddish brown, all parts K-. Conidia {microconidia) cylindrical,<br />
4-6x0-8-1 /xm.<br />
Chemistry: Thallus K— , C— , PD- ; apo<strong>the</strong>cia sections C— ; no substances detected by t.l.c.<br />
Observations: Micarea muhrii is characterized by its convex-adnate, never tuberculate,<br />
usually dark grey apo<strong>the</strong>cia, greenish upper hymenium, dark reddish brown hypo<strong>the</strong>cium<br />
(without green or purple tinges, even in K), and simple spores. It recalls <strong>the</strong> mainly saxicolous<br />
M. lutulata in its pigmentation, but that species differs in having convex-subglobose, <strong>of</strong>ten<br />
tuberculate apo<strong>the</strong>cia, smaller spores, and a larger celled phycobiont. The lignicolous M.<br />
melaeniza is ano<strong>the</strong>r species with similar pigmentation, but has smaller, markedly convex to<br />
tuberculate apo<strong>the</strong>cia, smaller spores, and sessile or stalked pycnidia.<br />
Habitat and distribution: Often in abundance on lignum <strong>of</strong> decorticate logs that lie across<br />
streams and probably become inundated at times; collected once on a periodically inundated<br />
boulder in a stream. So far, known only from Varmland in Sweden, from where it has been<br />
found in at least four localities by Lars-Erik Muhr, in whose honour I have <strong>the</strong> pleasure <strong>of</strong><br />
naming this species.<br />
28. Micarea myriocarpa V. Wirth & Vezda ex Coppins, sp. nov.<br />
(Figs 23C, 47C; Map 14)<br />
Micarea myriocarpa V. Wirth & Vezda in Poelt & Vezda, Bestimmungsschl. europ. Flechten, Erganzungsheft<br />
I: 161 (1977); nom. nudum (Art. 32). - Micarea myriocarpa V. Wirth & Vezda in V. Wirth,<br />
Flechtenfl.: 341, 345 (1980); nom. nudum (Art. 32).<br />
Thallus effusus, farinoso-granulosus, tenuissimus vel crassiusculus ad 0-3 mm crassus, pallide viridis vel<br />
viridulo-fulvus. Algae cellulis ± globisis, c. 4-1 ^tm diam, conglomeratis in granula goniocystiformes.<br />
Apo<strong>the</strong>cia immarginata, primum convexo-hemisphaerica mox tuberculata, pallide vel obscure rufa, vel<br />
fusca, 0-1-0-25 mm diam. Hymenium 25-35 /xm altum, ± hyalinum vel dilute rufo-brunneolum, cum vittis<br />
verticalibus obscure rufo-brunneolis, K — . Ascosporae oblongae vel oblongo-ovoideae, rectae vel ±<br />
curvatae, 5-5-8-5 x 1 •5-2-5 /xm. Paraphyses aliquantum paucae, dimorphae:p.p. hyalinae, laxae, simplices<br />
vel parce ramosae, interdum anastomosantes, graciles, 0-8-1-2 fxm latae, apicibus interdum incrassatis ad<br />
1-8 fim \ahs; p.p. pigmentiferae, fasciculatae, plerumque simplices, crassae, 2-3 /xm latae. Hypo<strong>the</strong>cium<br />
rufo-fuscum vel armeniaco-fuscum, K — , HNO3<br />
- . Excipulum nullum. Pycnidia pauca et inconspicuosa, ±
162 BRIAN JOHN COPPINS<br />
sessilia, doliiformia, rufo-fusca, c. 25-30 /tm diam. Conidia breviter cylindrica, 2-5-3-2X 1-1-3 /u.m. Thallus<br />
et apo<strong>the</strong>cia K— , C—<br />
, PD<br />
— ; sine materia chemica.<br />
Typus: Germania: 'Baden, Nordschwarzwald, Schurmsee bei Schonmiinzach bei Schwarzenberg, alt.<br />
790 m, Tannen am Seeausfluss', 25 v 1976, leg. V. Wirth (hb V. Wirth 6085 - holotypus!).<br />
Thallus effuse, pale green or greenish buff, scurfy farinose-granular, thin or thickish up to 0-3<br />
mm thick. Phycobiont ?micareoid, grouped into small goniocyst-like clusters c. 10-15 ^tm diam;<br />
cells ± globose 4-7 ixxn diam, or ellipsoid 5-7x3-5-5 /xm.<br />
Apo<strong>the</strong>cia numerous, immarginate, at first convex-hemispherical, soon becoming ± globose,<br />
<strong>of</strong>ten becoming tuberculate, pale to dark reddish brown, rarely brown-black, 0- 1-0- 15 mm<br />
diam, or to 0-25 mm diam when tuberculate. Hymenium c. 25-35 ^x.m tall, hyaline or dilute<br />
brownish, with dark brown, vertical streaks due to fasciculate pigmented paraphyses; K-,<br />
HNO3— . Asci cylindrical-clavate, 21-25x5-7 jxm. Spores oblong or oblong-ovoid, straight or<br />
slightly curved, simple or 1 -septate, 5 •5-8-5x1 •5-2-5 ^cm. Paraphyses ra<strong>the</strong>r scanty, <strong>of</strong> two<br />
types: p.p. hyaline throughout, evenly distributed, simple or sparingly branched (especially<br />
above), sometimes anastomosing, thin, 0^8-l-2 /otm wide, sometimes widening towards <strong>the</strong>ir<br />
apices to 1-8 /xm; p.p. pigmented ± throughout, in small fasciculate clusters arising from <strong>the</strong><br />
hypo<strong>the</strong>cium, stout, 2-3 /am wide. Hypo<strong>the</strong>cium 35-85 jxm tall, reddish brown or dull orange-<br />
brown, K-, HNO3— ; hyphae coated with dense brownish pigment and c. 2-3 /xm wide,<br />
interwoven but becoming vertically orientated towards <strong>the</strong> hymenium and sometimes continuing<br />
into it as pigmented, fasciculate paraphyses; ascogenous hyphae similarly pigmented but<br />
with short swollen cells, to 4 /xm wide. Excipulum not evident, even in sections <strong>of</strong> young<br />
apo<strong>the</strong>cia.<br />
Map 14 Micarea myriocarpa # 1950 onwards O Before 1950
LICHEN GENUS MICAREA IN EUROPE 163<br />
Pycnidia occasionally present but very inconspicuous, ± sessile, doliiform reddish brown, c.<br />
25-35 ixm tall and 25-30 /xm diam; wall reddish brown or dull orange-brown, K— , HNO3—<br />
Conidiogenous cells ± cylindrical to ampulliform, 4-5 x 1-1 -2 /xm but base sometimes swollen to<br />
2 jxm wide. Conidia (mesoconidia) short cylindrical, sometimes faintly biguttulate, 2-5-3-2xl-<br />
1-3 fxm.<br />
Chemistry: Thallus K— , C— , PD— ; apo<strong>the</strong>cia sections C— ; no substances detected by t.l.c.<br />
Observations: Micarea myriocarpa is characterized by its numerous, small, ± globose to<br />
tuberculate, reddish brown apo<strong>the</strong>cia, darkish (but never blackish, even in thick sections)<br />
hypo<strong>the</strong>cium, and small, narrow, 0-1-septate spores. The apo<strong>the</strong>cial tissues completely lack any<br />
greenish pigmentation, unlike in M. tuberculata and Psilolechia clavulifera, species <strong>of</strong> similar<br />
habitats with which M. myriocarpa has been confused. M. botryoides is ano<strong>the</strong>r common<br />
associate <strong>of</strong> M. myriocarpa, but can be distinguished by its black apo<strong>the</strong>cia, darker hypo<strong>the</strong>cium,<br />
larger spores, and numerous stalked, black pycnidia.<br />
Habitat and distribution: M. myriocarpa is faithful to <strong>the</strong> Micareetum sylvicolae and is found<br />
especially on exposed roots in dry underhangs under trees on steep slopes in valley woodlands.<br />
In <strong>the</strong> same situations it has been collected also on loose stones and mats <strong>of</strong> dry bryophytes. An<br />
additional habitat for M. myriocarpa is <strong>the</strong> dry undersides <strong>of</strong> stumps (especially conifers),<br />
situations which are microclimatically identical to rocky underhangs. In Britain, M. myriocarpa<br />
is widely distributed, but little collected, in western and nor<strong>the</strong>rn districts. On <strong>the</strong> Continent <strong>the</strong><br />
species is little known, but I have seen material from western Norway as far north as Trondheim,<br />
and Varmland in Sweden; and Wirth {loc. cit.) records it from <strong>the</strong> nor<strong>the</strong>rn Schwarzwald in<br />
Germany (<strong>the</strong> type locality).<br />
29. Micarea nigella Coppins, sp. nov.<br />
(Figs24A,47B;Mapl5)<br />
Thallus effusus, immersus vel tenuissimus, albidus vel viridi-griseus. Algae cellulis 4-7 /xm diam.<br />
Apo<strong>the</strong>cia primum subglobosa mox tuberculata, immarginata, atra, 0- 1-0-3 mm diam. Hymenium 25-30<br />
/Ltm altum, ± incoloratum, autem parte summa (epi<strong>the</strong>cio) purpureo-fusca K-l- olivacea. Ascosporae<br />
ellipsoideae, ovoideae vel oblongo-ovoideae, simplices 6-5-11 x2-5-4 /xm. Paraphyses aliquantum<br />
paucae, dimorphae: p.p. hyalinae, laxae, ramosae, gracillimae, 0-7-1 /u,m latae, apicibus vulgo leviter<br />
incrassatis ad 2 /xm latis; p.p. pigmentiferae, fasciculatae, plerumque simplices, crassae, 2-2-5 ^tm latae.<br />
Hypo<strong>the</strong>cium obscure purpureo-fuscum, K+ obscure olivaceum. Excipulum paulum evolutum mox<br />
reflexum. Pycnidia numerosa, atra, sessilia vel stipitata, interdum ramosa, c. 40-300 )u,m alta et 40-80 /xm<br />
lata, parietibus purpureo-fuscis, K-l- obscure olivaceis. Conidia ellipsoidea ad breviter cylindrica, 3-4-<br />
4-3x1-2-1-6 ixm. Thallus et apo<strong>the</strong>cia K-, C-, PD-.<br />
Typus: Dania, Jyllandia, c. 16 km septentriones e Hobro, Rold Skov, Torstedlund Skov, ad truncum<br />
decorticatum vetustum coniferarum in sylva mixta, 8 viii 1979, B. J. Coppins 4429 (E - holotypus).<br />
Thallus effuse, endoxylic, or ± epixyhc as a thin whitish to pale green-grey crust, but not<br />
forming discrete granules or areolae. Phycobiont micareoid, cells 4-7 pm diam.<br />
Apo<strong>the</strong>cia few to numerous, immarginate, subglobose at first, <strong>of</strong>ten becoming tuberculate,<br />
black, matt, 0-1-0-2 pm diam, or to 0-3 mm diam when tuberculate. Hymenium 25-30 ^im tall,<br />
hyaline or tinged dull purplish brown in places, <strong>of</strong>ten with darker, purplish brown vertical<br />
streaks; upper part (2-15 pm) irregularly pigmented purplish brown; K-h dull green, HNO3-Ired.<br />
Asci clavate, 22-27x10-11 pm. Spores ellipsoid, ovoid or oblong-ovoid, simple, 6-5-<br />
12x2-5-4 /xm. Paraphyses scanty, <strong>of</strong> two types: p.p. evenly distributed, branched, hyaUne, thin,<br />
0-7-1 pm wide, with apices sometimes widening to 2 ^tm but not becoming pigmented although<br />
embedded in pigmented gel-matrix; p.p. scattered or in small fascicles, mostly simple, stout, 2-3<br />
pm wide, coated ± throughout by dark pigment. Hypo<strong>the</strong>cium 70-100(-160) pm tall, mottled,<br />
dark purplish brown, K-l- dark dull green, HNO3-I- purple-red; hyphae coated with densepigment<br />
and c. 1-5-3 pm wide, interwoven but becoming vertically orientated towards <strong>the</strong><br />
hymenium and sometimes continuing into it as stout, pigmented paraphyses; ascogenous<br />
hyphae similarly pigmented but with short, swollen cells, to 4 pm wide. Excipulum indistinct-<br />
.
164 BRIAN JOHN COPPINS<br />
sometimes evident in young apo<strong>the</strong>cia as a dark (concolorous with hypo<strong>the</strong>cium) narrow zone<br />
forming an edge to <strong>the</strong> reflexed part <strong>of</strong> <strong>the</strong> hymenium; hyphae radiating, branched and<br />
anastomosing, c. 0-8-1 -5 /xm wide.<br />
Pycnidia numerous, black, sessile or more usually stalked, 60-300 jxm tall (including stalk)<br />
and 40-80 jxm diam; stalks (pycnidiophores) simple or sometimes branched bearing up to 4<br />
pycnidia; stalk and pycnidial wall tissues dark purplish brown, K+ dark green, HNO3+<br />
purple-red. Conidiogenous cells A-A-1-6 /xm tall, with a cylindrical neck 1 •9-3-7 fxm tall and<br />
0-8-1-4 fxm wide, and with a swollen base 2-4-3(-3-7) /xm wide, <strong>the</strong> wall <strong>of</strong> which is <strong>of</strong>ten<br />
pigmented; percurrent proliferations not seen. Conidia (mesoconidia) ellipsoid or short cylin-<br />
drical, sometimes faintly biguttulate, 3-4-4-3x1-2-1-6 /xm.<br />
Chemistry: Thallus K- , C- , PD- ; apo<strong>the</strong>cia sections C- ; no substances detected by t.l.c.<br />
Observations: Micarea nigella belongs to <strong>the</strong> group <strong>of</strong> lignicolous species (including M.<br />
contexta, M. eximia, M. melaeniza, M. misella, M. olivacea, and M. rhabdogena) with an<br />
endoxylic or indistinct thallus and small, ± globose to tuberculate, black apo<strong>the</strong>cia. It is<br />
characterized by <strong>the</strong> purple-brown, K-h green pigment in <strong>the</strong> hymenium, hypo<strong>the</strong>cium and<br />
pycnidial tissues, simple spores and stalked pycnidia. It is most likely to be confused in <strong>the</strong> field<br />
with o<strong>the</strong>r species with stalked, black pycnidia, namely M. melaeniza and M. misella. M.<br />
melaeniza differs in having a bright green hymenium, a red-brown hypo<strong>the</strong>cium that does not<br />
turn green in K, a different pycnidial pigmentation and shorter mesoconidia. M. misella has a<br />
K-l- violet hymenium, a ± hyaline hypo<strong>the</strong>cium and an olivaceous, K+ violet pigment in <strong>the</strong><br />
pycnidial wall tissues. Apart from <strong>the</strong>ir lack <strong>of</strong> stalked pycnidia, M. contexta, M. eximia, and M.<br />
Map 15 Micarea nigella # -I-<br />
Micarea olivacea ®<br />
J
LICHEN GENUS MICAREA IN EUROPE 165<br />
olivacea can be distinguished from M. nigella by <strong>the</strong>ir 1-septate spores. M. rhabdogena has a<br />
brown (K+ dissolving) pigment in <strong>the</strong> upper hymenium, and smaller spores. M. nigella could be<br />
confused with diminutive lignicolous forms <strong>of</strong> M. melaena, but that species usually has a bright<br />
green hymenium, larger and 1-3-septate spores, and never has stalked pycnidia.<br />
Habitat and distribution: M. nigella occurs on ra<strong>the</strong>r s<strong>of</strong>t lignum <strong>of</strong> stumps <strong>of</strong> conifers and<br />
birch, in deeply shaded situations. Primarily, it appears to be an inhabitant <strong>of</strong> conifer<br />
woodlands, and has been found both in native pine forest (Black Wood <strong>of</strong> Rannoch) and mature<br />
conifer plantations. It is known from only three localities: north Jylland in Denmark, <strong>the</strong><br />
sou<strong>the</strong>rn central highlands <strong>of</strong> Scotland and north-east England; and, although undoubtedly a<br />
ra<strong>the</strong>r rare species, it has surely been much overlooked, at least in north-west Europe.<br />
30. Micarea nitschkeana (Lahm ex Rabenh.) Harm.<br />
(Fig. 24B;Mapl6)<br />
in Bull. Soc. Sci. Nancy II, 33: 64 (1899). - Bilimbia nitschkeana Lahm ex Rabenh., Lich. Europ. Exs.<br />
583 (1861). - Micarea denigrata var. nitschkeana (Lahm ex Rabenh.) Hedl. in Bih. K. svenska<br />
VetenskAkad. Handl. Ill, 18 (3): 79, 90 (1892). - Bacidia nitschkeana (Lahm ex Rabenh.) Zahlbr. in<br />
Annln naturh. Mas. Wien 22: 342 (1905). Type: Germany, Nordrhein-Westfalen, between Miinster and<br />
Nobriskrug, on Pinus in wood, T. R. J. Nitschke, Rabenh. Lich. Eur. Exs. 583 (M - lectotype!;<br />
isolectotypes: BM!, BM ex K!, M!).<br />
Lecidea spododes Nyl. in Flora, Jena 52: 410 (1869). - Bacidia spododes (Nyl.) Zahlbr. , Cat. lich. univ. 4:<br />
151 (1926). Type: England, Hampshire, Lyndhurst, New Forest, old pales, /. M. Crombie (H-NYL<br />
18819- lectotype!; isolectotypes: BM!, H-NYL 18820!).<br />
Bacidia nitschkeana f. microcarpa Erichsen in Verh. hot. Ver. Prov. Brandenb. 71: 97 (1929). Type: West<br />
Germany, Schleswig-Holstein, Eckernforde, near Levensau, Felmerholz, on Picea twigs, 9 xi 1924,<br />
C. F. E. Erichsen (HBG-holotype!).<br />
? Bilimbia spododes f. fusca B. de Lesd., Rech. Lich. Dunkerque: 200 (1910). Type: France, Nord,<br />
Dunkerque, Malo-Terminus, on dead branch <strong>of</strong> Hippophae, B. de Lesdain (not seen).<br />
?Bilimbia spododes f. livida B. de Lesd., Rech. Lich. Dunkerque, Suppl. I: 120 (1914). Type: France,<br />
Nord, Dunkerque, Zuydcoote, on dead branch oi Hippophae, B. de Lesdain (not seen).<br />
? Bilimbia spododes var. nigra B. de Lesd., Rech. Lich. Dunkerque, Suppl. I: 120 (1914). Type: France,<br />
Nord, Dunkerque, Malo-Terminus, dunes, on dried rhizome oiAmmophila arenaria, B. de Lesdain (not<br />
seen).<br />
Thallus effuse, usually forming small patches but sometimes wide-spreading, sometimes<br />
partly endoxylic but usually developed on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> substratum as crowded, <strong>of</strong>ten<br />
contiguous, convex to subglobose areolae. Areolae especially well developed around <strong>the</strong><br />
apo<strong>the</strong>cia, dull greenish white to green-grey, surface matt, c. 40-200 /xm diam; in section,<br />
without a well defined cortex and not surrounded by an amorphous covering layer, outermost<br />
hyphae sometimes surrounded by dilute olivaceous, K-(- violet pigment. Thallus sometimes<br />
blackish and scurfy due to disruption by invading dematiaceous fungi and non-lichenized algae.<br />
Phycobiont micareoid, cells 4-7 /am diam.<br />
Apo<strong>the</strong>cia scattered or, more usually, numerous and crowded, frequently contiguous or<br />
confluent, adnate, plane to convex-hemispherical, sometimes becoming tuberculate, immarginate<br />
or occasionally (especially when young) with an indistinct margin that is flush with <strong>the</strong> level<br />
<strong>of</strong> <strong>the</strong> disc, grey-black to black, or rarely whitish to pale grey-brown (shade forms), sometimes<br />
paler at <strong>the</strong> margin, matt. c. 0- 1-0-3 mm diam, or to 0-4 mm diam when tuberculate. Hymenium<br />
30-40 /xm tall, upper part dilute olivaceous to olivaceous, K+ violet, HNO34- red, C+ violet,<br />
also C-l- orange-red throughout due to gyrophoric acid; olivaceous pigment mostly confined to<br />
<strong>the</strong> gel-matrix, only rarely (in old, dark apo<strong>the</strong>cia) closely bound to <strong>the</strong> walls <strong>of</strong> <strong>the</strong> paraphyses.<br />
Asci clavate 25-40x9-5-11 jxm. Spores fusiform, mostly curved, (l-)3(-4)-septate, 10-17<br />
(-19)x2-5-3(-3-5) pun. Paraphyses numerous, branched and anastomosing l-l-5(-l-7) jxm;<br />
apices not or only slightly incrassate, and mostly without closely adhering pigment. Hypo<strong>the</strong>cium<br />
30-50 /am tall, hyaline; hyphae interwoven, c. 1-1-5 /am wide; ascogenous hyphae with<br />
swollen cells c. 2-5 /am wide. Excipulum distinct in sections <strong>of</strong> young, ± plane apo<strong>the</strong>cia, but
166 BRIAN JOHN COPPINS<br />
reflexed and obscured in older, more convex apo<strong>the</strong>cia; hyaline or dilute olivaceous (K+ violet)<br />
at outer edge; hyphae radiating, branched and anastomosing, c. 1-1-5 /jlui wide.<br />
Pycnidia indistinct but usually present, immersed in areolae; walls hyaline throughout, or<br />
olivaceous (K+ violet) around <strong>the</strong> ostiole; <strong>of</strong> three types: (a) c. 60-80 /xm diam, ostiole<br />
sometimes widely gaping; Conidia {macroconidia) curved or hamate, 1-3-septate, 12-26 xc. 1<br />
/Lim; (b) c. 50-100 (xm diam, ostiole <strong>of</strong>ten gaping and conidia extruded as a white blob; conidia<br />
(mesoconidia) short-cylindrical or obovate-oblong, apices rounded but <strong>of</strong>ten distinctly truncate<br />
at proximal end, sometimes biguttulate, (3-)3-5-5(5-7)xl-l-5 /xm; (c) c. 40-50 /xm diam,<br />
ostiole never gaping; conidia (microconidia) narrowly fusiform or bacilliform, (4-7-)5-5-<br />
7-5x0-7-0-8/>tm.<br />
Chemistry: Thallus K-, PD-; sections <strong>of</strong> thallus and apo<strong>the</strong>cia C+ orange-red (also, parts<br />
with olivaceous pigment, C+ violet); t.l.c: gyrophoric acid. If thallus is heavily parasitized and<br />
scurfy <strong>the</strong> C+ orange-red reaction may be difficult to obtain and gyrophoric acid may not be<br />
detectable by t.l.c.<br />
Observations: M. nitschkeana is closely related, and similar in most respects, to M. denigrata<br />
{q.v.) with mainly 1-septate spores, and M. globulosella (q.v.) with ± acicular or rod-shaped<br />
0-3 (-6) -septate spores. Like M. denigrata, M. nitschkeana has three pycnidial anamorphs;<br />
usually only one or two <strong>of</strong> <strong>the</strong>se states are found on any given thallus, but all three states (plus<br />
apo<strong>the</strong>cia) have been noted on a single specimen from Scotland {Coppins 3369). In <strong>the</strong> field M.<br />
nitschkeana can be confused with species such as M. cinerea, M. lignaria, and M. peliocarpa, but<br />
<strong>the</strong>se species usually have larger apo<strong>the</strong>cia and areolae, and are quite different when examined<br />
microscopically.<br />
Habitat and distribution: M. nitschkeana is widely distributed in <strong>the</strong> <strong>British</strong> Isles at low<br />
altitudes (below 300 m), and is most frequently found on twigs or small branches <strong>of</strong> various<br />
deciduous and coniferous trees, e.g. Acer, Aesculus, Alnus, Betula, Fraxinus, Prunus padus,<br />
Quercus, Salix, Sambucus, Larix, and Picea; or on <strong>the</strong> old or dead stems <strong>of</strong> smaller shrubby<br />
plants, especially Calluna and Ulex, but also Erica cinerea, Myrica, Rosa, Sarothamnus,<br />
Viburnum, and Juniperus. It has been found to be abundant on old stems and Utter <strong>of</strong> Calluna in<br />
some lowland heaths in eastern England (Coppins & Shimwell, 1971), Scotland and Denmark,<br />
where it is commonly associated with Lecanora conizaeoides and Scoliciosporum chlorococum;<br />
such communities are referable to <strong>the</strong> Bacidietum chlorococcae . O<strong>the</strong>r associated species on<br />
twigs and small branches include Lecanora pulicaris, L. symmicta agg., L. icmalea, Mycoblastus<br />
sterilis, Fuscidea lightfootii (western districts), Micarea prasina, Graphis elegans, Stenocybe<br />
pullatula (on Alnus) and small thalli <strong>of</strong> Hypogymnia physodes, H. tubulosa, Parmelia subaurifera,<br />
P. sulcata, Platismatia glauca, Evernia prunastri, and Ramalina fairinacea. It has<br />
occasionally been met with on lignum <strong>of</strong> fallen conifers (Larix and Pinus) or fence posts, habitats<br />
more characteristic <strong>of</strong> M. denigrata. Associated species on lignum include Lecanora confusa, L.<br />
symmicta agg. , Lecidea granulosa agg., L. icmalea, Mycoblastus sterilis, Hypogymnia physodes,<br />
Parmelia saxatilis, and Platismatia glauca. Occurrences on rock are rare, but a few collections<br />
have been made on sandstone or flint stones in heathland, with Lecidea granulosa agg., L.<br />
icmalea, and Micarea melaena as associates. It has also been found toge<strong>the</strong>r with Scoliciosporum<br />
umbrinum on a plastic carton in a heathland in <strong>the</strong> Isle <strong>of</strong> Wight (Brightman & Seaward, 1977).<br />
M. nitschkeana is widely distributed through much <strong>of</strong> nor<strong>the</strong>rn Europe. In Scandinavia it has a<br />
sou<strong>the</strong>rn distribution and I have not seen material <strong>of</strong> it from north <strong>of</strong> latitude 62°N. It usually<br />
occurs in lowland situations, but it attains higher altitudes in <strong>the</strong> Alps (920 m) and <strong>the</strong> Nizke<br />
Tatry <strong>of</strong> Czechoslovakia (1250 m). Additional phorophytes from extra-<strong>British</strong> material include<br />
Hippophae (France: dunes near Calais), Vaccinium uliginosum (Germany: Schleswig-<br />
Holstein), and Pinus mugo subsp. pumilio (heathlands <strong>of</strong> sou<strong>the</strong>rn Germany). From outside<br />
Europe I have seen only one specimen: USA, nor<strong>the</strong>rn California (near San Francisco), where it<br />
occurred on fence ppsts.<br />
Exsiccata: Arnold Lick. Exs. Ill (BM ex K, M), 503 A, B (H-NYL, M); 503 C (BM ex K, H-NYL, M),<br />
503 D (H-NYL, M). Arnold Lick. Mon. 48 (BM ex K, M); 270 (BM ex K). Britz. Lick. Exs. 829 (M).
LICHEN GENUS MICAREA IN EUROPE 167<br />
Map 16 Micarea nitschkeana # 1950 onwards O Before 1950<br />
Claudel & Harm. Lich. Gall. 89 (L, O). Harm. Lich. Loth. 853 (M). Hepp Flecht. Eur. 20 (BM, E, L, M),<br />
llp.p. (E). Krypt. Exs. Vindob. 1232 (M). Lojka Lich. Univ. 137 (BM ex K, M). Magnusson Lich. Sel.<br />
Scand. 340 (M). Malbr. Lich. Norm. 287 (M). Malme Lich. Suec. 25 (M, S). Rabenh. Lich. Eur. 582, 583<br />
(BM, BM ex K, M). Rasanen Lich. Fenn. 642 (M). Zahlbr. Lich. Rar. 110 (BM). Zwackh Lich. Exs. 470<br />
['bei Miinster in Westfalen'] (H-NYL 18825), 470 bis ['Bei Delbrueck in Kreise Paderborn.'] (H-NYL<br />
18822, M), 534 (H-NYL, M), 587 (H-NYL, M).<br />
31. Micarea olivacea Coppins, sp. nov.<br />
(Figs 24C, 47A; Map 15)<br />
Thallus effusus, endoxylicus vel epixylicus aut epilithicus, tenuissimus, inaequalis, interdum leviter<br />
areolatus, albidus vel viridio-griseus. Algae cellulis 4-7 ^tm diam. Apo<strong>the</strong>cia immarginata, primum<br />
convexa vel subglobosa mox tuberculata, atra, 0- 1-0-3 mm diam, aut ad 0-4 mm diam ubi tuberculata.<br />
Hymenium 30-35 />tm altum, dilute sordide olivaceum cum vittis verticalibus atro-olivaceis, K-l- clare<br />
olivaceo-virescens. Ascosporae oblongae vel ovoideo-oblongae, rectae, (O-)l-septatae, (7-)9-12-3x2-5-<br />
3-5 jxm. Paraphyses numerosae, dimorphae: p.p. hyalinae, laxae, ramosae et anastomosantes, graciles, c.<br />
1-1-2 /xm latae, apicibus interdum leviter incrassatis ad 2 fxm latis; p.p. pigmentiferae, fasciculatae,<br />
plerumque simplices, crassae, 2-3 ^tm latae. Hypo<strong>the</strong>cium sordide atro-olivaceum vel olivaceo-fuscum,<br />
K-l- virescens. Excipulum paulum evolutum mox reflexum. Pycnidia numerosa sed inconspicua, ±<br />
immersa, 25-50 ^tm diam., parietibus sordide olivaceis vel olivaceo-fuscis, K-l- clare olivaceis. Conidia<br />
brevitercylindrica, 3-4-2x1-1-3 />tm. Thallus ei apo<strong>the</strong>cia K- ,C-, PD-.<br />
Typus: Caledonia, Mull, Aros, Druimfin, in ligno duro, 15 v 1968, P. W. James (BM - holotypus).
168 BRIAN JOHN COPPINS<br />
Thallus effuse, endoxylic to epixylic, or epilithic, forming a thin, whitish grey or greenish grey,<br />
uneven, sometimes weakly areolate crust; <strong>of</strong>ten appearing scurfy due to invasion by foreign<br />
fungi and algae. Phycobiont micareoid, cells 4-7 /xm diam.<br />
Apo<strong>the</strong>cia numerous, immarginate, convex to subglobose, <strong>of</strong>ten tuberculate, black, matt or<br />
slightly glossy, surface minutely roughened, 0- 1-0-25 mm diam, or to 0-4 mm diam when<br />
tuberculate. Hymenium 30-35 /xm tall, without a distinct upper part (epi<strong>the</strong>cium), dilute sordid<br />
olivaceous with dark olivaceous vertical streaks, K+ green intensifying, HNO3+ red. Asci<br />
clavate, 35-33x9-5-11 /xm. Spores oblong or ovoid-oblong, straight, (O-)l-septate, (7-)<br />
9-12-3x2-5-3-5 />im. Paraphyses numerous, <strong>of</strong> two types: p.p. hyaline throughout, evenly<br />
distributed, branched and <strong>of</strong>ten anastomosing, ra<strong>the</strong>r thin, 1-1-2 /^m wide, apices sometimes<br />
widening to 2 /am and sometimes with colourless, oily, refractive contents; p.p. broad, 2-3 /xm<br />
wide, mostly simple, grouped in small fascicles and <strong>of</strong>ten embedded in dense pigment.<br />
Hypo<strong>the</strong>cium 40-70 /xm tall, dark sordid olivaceous or olive-brown, K+ green intensifying,<br />
HNO3+ red; hyphae embedded in greenish gel-matrix but walls not deeply pigmented, c. 1-5-2<br />
/am wide, interwoven but becoming vertically orientated towards <strong>the</strong> hymenium and sometimes<br />
continuing in to it as fasciculate paraphyses; ascogenous hyphae with short swollen cells, to 5 /am<br />
wide. Excipulum indistinct and soon reflexed, sometimes evident in sections <strong>of</strong> young apo<strong>the</strong>cia<br />
as a hyaline or dilute sordid olivaceous, non-amyloid zone c. 20 /am wide; composed <strong>of</strong> radiating,<br />
much branched and anastomosing hyphae 0-8-1-5 /am wide.<br />
Pycnidia numerous but inconspicuous, ± immersed, 25-50 /am diam; walls sordid olivaceous<br />
or olive-brown, K-l- green intensifying, HNO3-I- red. Conidiogenous cells elongate-ampulliform<br />
to subcyUndric, 4—5-5x1-5-3 /am. Conidia (mesoconidia) short cylindrical, sometimes faintly<br />
biguttulate, 3-4-2x1-1-3 /am.<br />
Chemistry: Thallus K— , C— , PD— ; apo<strong>the</strong>cia sections C— ; no substances detected by t.l.c.<br />
Observations: Micarea olivacea differs from M. eximia in its less brightly coloured hymenium<br />
pigmentation that is not concentrated in <strong>the</strong> upper part, more numerous and broader paraphyses,<br />
shorter and slightly broader spores with rounded apices, and shorter mesoconidia. From<br />
M. nigella it can be distinguished by <strong>the</strong> complete absence <strong>of</strong> purple pigmentation in water<br />
mounts, more numerous and broader paraphyses, more elongate and 1-septate spores, and ±<br />
immersed (never stalked) pycnidia. M. olivacea is easily confused with epruinose forms <strong>of</strong><br />
<strong>the</strong> common lignicole Lecidea turgidula Fr., but that species has a dark green or olive-brown<br />
excipulum <strong>of</strong> conglutinated hyphae that do not separate in K, a paler hypo<strong>the</strong>cium, and a thick<br />
walled, large celled phycobiont with cells 12-16 /am diam. In addition, <strong>the</strong> conidia <strong>of</strong> L. turgidula<br />
are smaller, c. 3-3-5x1-5-1-8 /am. When on rock M. olivacea can be confused with M.<br />
tuberculata, but <strong>the</strong> latter has a more brightly coloured hymenium and hypo<strong>the</strong>cium, somewhat<br />
smaller spores, narrower asci, and a non-micareoid phycobiont with cells c. 5-10 /am when<br />
globose or up to 15 X 7 /am when ellipsoid . M. tuberculata has similar pycnidia and conidia , but its<br />
conidiogenous cells are more slender and about twice as long as those <strong>of</strong> M. olivacea.<br />
Habitat and distribution: M. olivacea is an apparently rare, but probably overlooked, species,<br />
being known from just two localities, both in Scotland. At <strong>the</strong> type locality it occurred on <strong>the</strong><br />
hard Ugnum <strong>of</strong> a stump by a conifer plantation; <strong>the</strong> collection contains no associate species. At<br />
<strong>the</strong> o<strong>the</strong>r locality it occurred with Rhizocarpon hochstetteri and Baeomyces rufus on shaded rock<br />
in a mature conifer plantation. Little more can be said <strong>of</strong> its ecology until it becomes better<br />
known, but I am inclined to believe that it is essentially a lignicolous species with a preference for<br />
hard lignum . If this is true <strong>the</strong>n care should be taken not to overlook it in <strong>the</strong> field for forms <strong>of</strong> <strong>the</strong><br />
common M. denigrata with a reduced thallus.
32. Micarea osloensis (Th. Fr.) Hedl.<br />
(Fig. 28B)<br />
LICHEN GENUS MICAREA IN EUROPE 169<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 83, 97 (1892). - Lecidea osloensis Th. Fr., Lich.<br />
Scand. 2: 524 (1874). Type: Norway, Oslo, summit <strong>of</strong> Ryenbjerget, on decaying plant debris, N. G. Moe<br />
(UPS-holotype! [t.l.c.: no substances]).<br />
Thallus effuse, indistinct, apparently <strong>of</strong> whitish grey, ± convex areolae, c. 50-70 /am diam, in<br />
<strong>the</strong> single specimen seen <strong>the</strong> thallus is obscured by a bleached non-lichenized alga, giving it a<br />
pale grey 'filmy' appearance. Phycobiont ^xohdhXy micareoid, cells 4-5-6-5 /am diam.<br />
Apo<strong>the</strong>cia numerous, ± evenly scattered, immarginate, convex to subglobose, black or<br />
brown-black, 0- 1-0-26 mm diam. Hymenium c. 30 /am tall; upper part (epi<strong>the</strong>cium) red-brown,<br />
K— , HNO3—<br />
or red tinge intensifying; remaining (lower) part dilute reddish brown with darker,<br />
red-brown, vertical streaks. Asci clavate, 26-30x 11-13 /am. Spores ellipsoid or ovoid-ellipsoid,<br />
simple, 6-9-5x3-4 /am. Paraphyses ra<strong>the</strong>r numerous, simple below, but mostly branched<br />
towards <strong>the</strong>ir apices, hyaline and 1-5-1-8 /am wide, but occasionally surrounded throughout<br />
<strong>the</strong>ir length by red-brown pigment and <strong>the</strong>n 1-7-2 /am wide; apices sometimes swollen to 3 /am<br />
wide, surrounded by an amorphous, red-brown, pigmented matrix that does not disperse in K.<br />
Hypo<strong>the</strong>cium c. 70-85 /am tall, dark reddish brown, K— , HNO3—<br />
Excipulum not evident, even in sections <strong>of</strong> young apo<strong>the</strong>cia.<br />
Pycnidia not seen.<br />
Chemistry: Thallus K— , C—<br />
detected by t.l.c.<br />
, PD—<br />
or red tinge intensifying.<br />
; sections <strong>of</strong> thallus and apo<strong>the</strong>cia C— ; no substances<br />
Observations: Micarea osloensis is characterized by its small, blackish apo<strong>the</strong>cia with internal<br />
reddish brown pigment that does not disperse in K, simple, ellipsoid spores, and terricolous<br />
habitat. In <strong>the</strong> field it is most likely to be overlooked for Lecidea uliginosa, but that species has a<br />
well developed, pseudoparenchymatous excipulum, paraphyses with a brown apical cap, larger<br />
spores, and 'Trapelioid' asci which do not have a deeply amyloid tholus.<br />
Habitat and distribution: Knowledge <strong>of</strong> M. osloensis is still restricted to its type ga<strong>the</strong>ring from<br />
Norway. The appearance <strong>of</strong> this material suggests that it occurred in a woodland clearing on <strong>the</strong><br />
site <strong>of</strong> an old bonfire (small amounts <strong>of</strong> ash are present), probably in a slight depression. The<br />
specimen is accompanied by <strong>the</strong> sterile thallus <strong>of</strong> Lecidea icmalea and a few plants <strong>of</strong> <strong>the</strong> moss<br />
Ceratodon purpureas. Dr H. Krog informs me that <strong>the</strong> hill Ryenberget is now well within <strong>the</strong> city<br />
limits <strong>of</strong> Oslo, and is now much affected by <strong>the</strong> results <strong>of</strong> urbanization, although not completely<br />
destroyed. It has an altitude <strong>of</strong> about 150-200 m, and was almost certainly covered in pine forest<br />
before it was engulfed by <strong>the</strong> city.<br />
33. Micarea peliocarpa (Anzi) Coppins & R. Sant.<br />
(Figs 3A-C, 25, 48, 53-54; Map 17)<br />
in Coppins & P. James in Lichenologist 11: 155 (1979). - Bilimbia peliocarpa Anzi in Atti Soc. ital. Sci.<br />
nat. 9: 250 (1866). - Bacidia peliocarpa (Anzi) Lettau in Hedwigia 52: 133 (1912). - Micarea violacea f.<br />
peliocarpa (Anzi) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 81, 92 (1892); nom. superfl.<br />
(Art. 63). Type: Italy, Piemonte, Novara, 'Sopra i muschi nei monti del Lago Maggiore (Locarno),<br />
scoperta dal padre capuccino Agostino Daldini' (? TO, not seen: request for loan not acknowledged); an<br />
unlocalized specimen labelled 'Bilimbia peliocarpa Anzi Neosymb. no. 44' [reference to original<br />
protologue] in Anzi's handwriting occurs in UPS(!) and is possibly an isotype.<br />
Biatora lignaria (3. [var.] conglomerata Hepp, Flecht. Eur., fasc. 5, no. 284 (1857). - Micarea violacea f.<br />
conglomerata (Hepp) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 81, 92 (1892). Type:<br />
Switzerland, 'bei Rifferschweil', on Finns bark, Hegetschweiler, Hepp Flecht. Eur. 284 (E - lectotype!;<br />
M-isolectotype!).<br />
Biatora lignaria var. saxigena Hepp, Flecht. Eur. fasc. 9, no. 510 (1860). Type: Germany, Hessen, 'auf<br />
Sandsteinfelsen bei Marburg', W. Uloth, Hepp Flecht. Eur. 510(E-lectotype!; M-isolectotype!). See<br />
note (i) below.
170 BRIAN JOHN COPPINS<br />
Bilimbia milUaria var. lignaria *[f.] livida Korber, Parerga Lich.: 171 (1860). Type: same as Biatora<br />
lignaria var. conglomerata Hepp.<br />
Lecidea sphaeroides var. leucococca Nyl. in Stizenb. in Nova Acta Leop. - Carol. 34 (2): 18, 1. 1 , f. D, 47-51<br />
(1867). Type: Finland, Tavastia australis, Evois ['Evo'], on lignum, 1865, J. P. Norrlin (H-NYL 18377lectotype!;<br />
H-isolectotype!).<br />
Bilimbia violacea Arnold in Flora, Jena 53: 473 (1870). - Lecidea violacea P. Crouan & H. Crouan ex Nyl.<br />
in Flora, Jena 45: 464 (1862), non Massal. (1852). - Micarea violacea (Arnold) Hedl. in Bih. K. svenska<br />
VetenskAkad. Handl. Ill, 18 (3): 80, 91 (1892). Type: France, Finistere, Brest, on schistose rock,<br />
Crouan (H-NYL 18716- isotype!). [Note: listed as 'Bacidia violacea (Crouan ex Nyl.) Arnold' by James<br />
(1965a); this should not be confused with Bacidia violacea (Arnold) Arnold, Flora, Jena 67: 581 (1884),<br />
which is a Bacidia s. str.]<br />
Bilimbia trisepta Hellb., Nerikes Lafflora: 11 (1871). - Biatora trisepta Naeg. ex Mull. Arg. in Mem. Soc.<br />
Phys. Hist. nat. Geneve 16: 403 (1862); as 'Naeg. mss. ex Dr Hepp', nom. inval. (Arts 32, 34). - Lecidea<br />
sabuletorum f. trisepta Stizenb. in Nova Acta Acad. Leop. -Carol. 34 (2): 47, t. 3, f. A, 35-62 (1867); nom.<br />
superfl. (Art. 63). - Bilimbia hypnophila b. [var.] trisepta Bausch in Ver. naturw. Ver. Karlsruhe 4: 127<br />
(1869); nom. superfl. (Art. 63). - Bilimbia sabuletorum d. [var.] trisepta Rabenh., Krypt.-Fl. Sachsen 2:<br />
187 (1870); nom. superfl. (Art. 63). - Bacidia trisepta (Hellb.) Zahlbr. in Engler & Prantl, Nat.<br />
Pflanzenfam. 1 (1*): 135 (1905). -Micarea trisepta (Hellb.) Wetmore in Pub. Mus. Mich. St. Univ. Biol.<br />
3: 284 (1968). Type: Germany, Hessen, 'auf Sandsteinfelsen bei Marburg', W. Uloth, Hepp Flecht. Eur.<br />
510 (E-neotype!; M-isoneotype!). See note (ii) below.<br />
Lecidea hemipolioides Nyl. in Flora, Jena 56: 294 (1873). - Bilimbia hemipolioides (Nyl.) A. L. Sm.,<br />
Monogr. Br. Lich. 2: 141 (1911). - Bacidia hemipolioides (Ny\.) Zahlbr. , Cat. lich. univ. 4: 114(1926).-<br />
Micarea violacea f. hemipolioides (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 80, 91<br />
(1892). Type: Channel Islands, Jersey, Rozel meadow, on rocks, 1873, C. Larbalestier (H-NYL 18713 -<br />
lectotype!; isolectotypes: H-NYL p.m. 4613!, BM!).<br />
Bilimbia naegelii f. lapiseda Th. Fr., Lich. Scand. 2: 379 (1874). Type: Sweden, Stockholm, Lyran, on<br />
rock, 1870, S. Almquist (IJ?S - lectotype!).<br />
Bilimbia milliaria f. livescens Th. Fr., Lich. Scand. 2: 383 (1874). Type: Sweden, Narke, Gotlunda, 'pa<br />
stenmuren mellan Stabacken och Hogby', on mosses, 1863, Blomberg (UPS - lectotype!).<br />
Lecidea albidolivens Nyl. in Flora, Jena 57: 10 (1874). - Bilimbia albidolivens (Nyl.) Blomb. & Forss.,<br />
Enum. PI. Scand.: 82 (1880). Type: Finland, Tavastia australis, Padasjoki, Nyystola, on lignum, 1872,<br />
E. A. Lang fVainioj (H-NYL 18775 - lectotype!; H - isolectotype!).<br />
Lecidea fraterculans Nyl. in Flora, Jena 58: 11 (1875). Type: Finland, Tavastia australis, Padasjoki,<br />
Nysstola, on rock, 1872, E. A. Lang [VainioJ (H-NYL 18704a - lectotype!; H-NYL 18704 - isolecto-<br />
type!).<br />
Lecidea triseptatula Nyl. in Flora, Jena, 58: 361 (1875). Type: Finland, Tavastia australis, HoUola, on<br />
lignum, 1874, E. A. Lang fVainioj (H-NYL 18709 -lectotype!; isolectotypes: H-NYL 18710!, H!).<br />
Bilimbia albicans Arnold in Flora, Jena 65: 140 (1882). Type: Germany, Bayern, Oberbayern, near Bad<br />
Tolz, W side <strong>of</strong> <strong>the</strong> Blombergs, on sandstone in wood, 5 ix 1880, Arnold, Lich. exs. 837 (BM -<br />
lectotype!).<br />
Micarea violacea f. cupreola Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 80, 91 (1892). Type:<br />
Sweden, Halsingland, Jarvso, on rock, viii 1891, Hedlund(S-\\o\oiypQ\).<br />
Lecidea triseptatuloides Harm, in Bull. Seanc. Soc. Sci. Nancy II, 33: 64 (1899). - Bacidia triseptatuloides<br />
(Harm.) Zahlbr., Cat. lich. univ. 4: 161 (1926). Type: France, Moselle, Bitche, on Pinus bark, xii 1893,<br />
Abbe Kieffer (ANGUC - holotype!).<br />
Micarea violacea f. exigua Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 81, 91 (1892); nom.<br />
inval. (Art. 26).<br />
Notes: (i) 'saxigena'. This epi<strong>the</strong>t has been applied at intra-specific levels to saxicolous forms <strong>of</strong> M.<br />
lignaria and M. peliocarpa. Its first valid publication was by Hepp (loc. cit.) as "Biatora lignaria (Ach.) var.<br />
saxigena (Leight) Hepp' with 'Lecidea milliaria (Fries) v. saxigena, Leight Lich. Brit. Exs. 210' given as a<br />
synonym. All examples seen by me <strong>of</strong> Leighton Lich. Brit. 210 (issued in 1856) are M. lignaria; whereas all<br />
examples <strong>of</strong> Hepp Flecht. Eur. 510 are M. peliocarpa. Hepp's illustration and description <strong>of</strong> spores are<br />
more applicable to M. peliocarpa than M. lignaria. The original valid introduction <strong>of</strong> <strong>the</strong> epi<strong>the</strong>t "saxigena'<br />
should, <strong>the</strong>refore, be typified on an example <strong>of</strong> Hepp Fl. Eur. 510 and attributed to Hepp alone.<br />
Theoretically "saxigena Leighton' could have been validated for saxicolous forms <strong>of</strong> M. lignaria by<br />
providing a description, excluding o<strong>the</strong>r names or specimens that would render it superfluous, and<br />
avoiding homonymy with previous combinations <strong>of</strong> "saxigena Hepp' ; but I have not found such a case in <strong>the</strong><br />
Hterature.<br />
(ii) "trisepta". This epi<strong>the</strong>t first appeared as "Biatora trisepta Naeg. mss. ex Dr Hepp' in <strong>the</strong> protologue <strong>of</strong>
LICHEN GENUS MICAREA IN EUROPE 171<br />
Patellaria salevensis Miill. Arg. (= Bacidia salevensis (Mull. Arg.) Zahlbr.). Miiller presumably regarded<br />
Naegeli's manuscript name as a synonym <strong>of</strong> his new species. B. salevensis (type in BM!) is a limestone<br />
species and is not a Micarea. As indicated by James (1965: 103) <strong>the</strong> first legitimate publication <strong>of</strong> <strong>the</strong> epi<strong>the</strong>t<br />
'trisepta' for a Bacidia-\\kt species is that by Hellbom {loc. cit). By including <strong>the</strong> epi<strong>the</strong>t in Bilimbia as '5.<br />
trisepta (Naeg.)', Hellbom was clearly using it in its traditional sense and not introducing it as an entirely<br />
new name. There is no indication that any <strong>of</strong> <strong>the</strong> authors using <strong>the</strong> epi<strong>the</strong>t 'trisepta Naeg.' had seen<br />
Naegeli's original material, and no such material has been found amongst Hepp's collections at BM.<br />
Because <strong>the</strong> original material is probably no longer in existence, neotypification is required. The epi<strong>the</strong>t<br />
'trisepta'' has been a long source <strong>of</strong> confusion, being used by various authors for several species, especially<br />
Micarea lignaria, M. melaena, M. nitschkeana, M. peliocarpa, and M. ternaria. However, it is clear from<br />
<strong>the</strong> descriptions and cited specimens in Stizenberger (loc. cit.), Bausch (loc. cit.), Hedlund (1892), and<br />
Vainio (1922) that <strong>the</strong> epi<strong>the</strong>t 'trisepta' has most commonly been applied to <strong>the</strong> species now called Micarea<br />
peliocarpa. Because 'trisepta' first appeared in <strong>the</strong> protologue <strong>of</strong> a saxicolous species <strong>the</strong> neotype chosen is<br />
an example <strong>of</strong> Hepp Flecht. Eur. 510 which was cited under, or in connection with, 'trisepta' in <strong>the</strong><br />
treatments <strong>of</strong> Stizenberger, Bauch, Hedlund, and Vainio.<br />
Thallus effuse, sometimes partly endocuticular, endophloeodal or endoxylic, more usually<br />
developed on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> substratum as rounded, shallow-convex, hemispherical, or ±<br />
globose areolae. Areolae scattered or more usually ± continguous, greenish white, or greywhite<br />
to blue-grey, occasionally becoming dark grey, <strong>of</strong>ten dark coloured on upper surface but<br />
pale and greenish white below, matt or slightly glossy, c. 40-200 ixm diam. Areolae in section,<br />
ecorticate but with an amorphous hyaline covering layer c. 2-5 /x,m thick; outermost hyphae<br />
<strong>of</strong>ten with grey-green to blue-green walls, K-, HNO3+ red. Phycobiont micareoid, cells 4-7<br />
/xm diam.<br />
Apo<strong>the</strong>cia scattered or more usually crowded and <strong>of</strong>ten contiguous, adnate, plane to convex,<br />
sometimes becoming tuberculate; sometimes with an indistinct margin that is flush with <strong>the</strong> level<br />
<strong>of</strong> <strong>the</strong> disc; rarely (as in shaded corticolous forms) whitish or ivory-white, usually pale lead-grey<br />
to grey-black, sometimes grey-brown, <strong>of</strong>ten whitish or paler at <strong>the</strong> margin; (0-12-)0- 14-0-4<br />
(-0-6) mm diam, or to 1 mm when tuberculate. Disc finely roughened, matt, or slightly glossy<br />
(when black); margin (when evident) smooth and <strong>of</strong>ten more glossy than <strong>the</strong> disc. Hymenium<br />
40-55 iJLm tall, usually olive-green or aeruginose (K— , HNO3-I- red) in upper part and ± hyaline<br />
below; in dark coloured apo<strong>the</strong>cia <strong>the</strong> green pigment <strong>of</strong>ten occurs as vertical streaks through <strong>the</strong><br />
hymenium. Asci clavate c. 40-55x12-17 /xm. Spores fusiform, clavate-fusiform, or oblongfusiform,<br />
<strong>of</strong>ten slightly curved, (l-)3(-5)-septate, (ll-)15-23(-24)x3-5(-6) /am. Paraphyses<br />
numerous, branched, <strong>of</strong>ten anastomosing, 1-1-5 /xm wide; apices <strong>of</strong>ten more richly branched<br />
and entangled, <strong>of</strong>ten slightly incrassate to c. 1-8 /xm, or to 2-5 /i,m due to thickening by green<br />
pigment. Hypo<strong>the</strong>ciumc. 40-70 />im tall, hyaUne or dilute straw; hyphae interwoven, c. 1-1 -7 /xm<br />
wide; ascogenous hyphae with swollen cells c. 2-4 /xm wide. Excipulum well developed, hyaline,<br />
or dilute straw in part, <strong>of</strong> richly branched and anastomosing, radiating hyphae c. 1-1-5 ^im.<br />
Pycnidia frequently present (especially on bark or lignum), <strong>of</strong> two types: (a) immersed in<br />
areolae, white or greenish around <strong>the</strong> ostiole, 140-200 /xm diam, ostiole <strong>of</strong>ten widely gaping;<br />
conidia (macroconidia) markedly curved and <strong>of</strong>ten sigmoid, <strong>of</strong>ten faintly 1-5-septate, 21-40(-<br />
50)x 1-1-5 fim; (b) ± sessile, white, 50-80 /xm diam, ostioles not, or only slightly, gaping;<br />
conidia (microconidia) narrowly fusiform-cylindrical, (5-)6-7(-7-7)x0-4-0-7 /xm.<br />
Chemistry: Thallus and whitish apo<strong>the</strong>cia K-, C-lorange-red;<br />
t.l.c: gyrophoric acid.<br />
red, PD— ; apo<strong>the</strong>cia in section C-l-<br />
Observations: M. peliocarpa occurs in a wide range <strong>of</strong> habitats and is concurrently variable,<br />
especially regarding <strong>the</strong> colour <strong>of</strong> its apo<strong>the</strong>cia and thallus. This variation is considered to be<br />
phenotypic, being due to <strong>the</strong> amount <strong>of</strong> green pigment produced in <strong>the</strong> hymenium and near <strong>the</strong><br />
surface <strong>of</strong> <strong>the</strong> thallus in response to environmental factors, particularly light. When on <strong>the</strong> bark<br />
<strong>of</strong> old trees in sheltered woodland <strong>the</strong> apo<strong>the</strong>cia are <strong>of</strong>ten very pale to blue-grey, more scattered<br />
than usual, and accompanied by numerous pycnidia; such forms are frequent in <strong>the</strong> ancient<br />
woodlands <strong>of</strong> <strong>the</strong> New Forest, Hampshire. There is some variation as to <strong>the</strong> production <strong>of</strong><br />
areolae, <strong>the</strong>se being <strong>of</strong>ten poorly developed or even absent when on cortex or lignum, and
172 BRIAN JOHN COPPINS<br />
usually abundant and well developed in more exposed situations on mossy rocks and peaty<br />
debris.<br />
M. peliocarpa is most closely related to M. alabastrites and M. cinerea (see under <strong>the</strong> former<br />
for fur<strong>the</strong>r discussion). Apart from <strong>the</strong>se two species, M. peliocarpa is most <strong>of</strong>ten confused with<br />
M. lignaria {q.v.) especially when occurring in exposed habitats where its apo<strong>the</strong>cia may be<br />
black and markedly convex. Similarly, confusion could occur with <strong>the</strong> rare M. ternaria {q.v.),<br />
which like M. peliocarpa has ra<strong>the</strong>r flattened apo<strong>the</strong>cia (in section) with a well developed<br />
excipulum. When fertile, M. leprosula has apo<strong>the</strong>cia very hke those <strong>of</strong> M. peliocarpa, but its<br />
thallus is composed <strong>of</strong> fragile areolae that readily dissolve into soredia and contains argopsin<br />
(PD+ red) as well as gyrophoric acid. In certain habitats, such as <strong>the</strong> stems <strong>of</strong> old shrubs (e.g.<br />
Calluna, Erica, and Ulex), M. peliocarpa could be mistaken for M. nitschkeana although <strong>the</strong><br />
latter has an olivaceous, K+ violet pigment in <strong>the</strong> hymenium, pycnidial walls and thallus, and<br />
smaller spores. Specimens collected from Xanthorion communities and attributed to M.<br />
peliocarpa (or one <strong>of</strong> its synonyms) are mostly referable to Bacidia naegelii, which has simple<br />
paraphyses with markedly swollen apices, an excipulum <strong>of</strong> coherent hyphae (in K), and a very<br />
different thallus structure.<br />
Habitat and distribution: M. peliocarpa is ra<strong>the</strong>r catholic in its choice <strong>of</strong> habitat. As an<br />
epiphyte on bark (or on bryophytes <strong>the</strong>reon) it is mostly commonly found on mature Quercus,<br />
but has also been collected from Alnus, Betula, Fagus, Fraxinus, Ilex, Salix, Larix, Pinus,<br />
Juniperus, Calluna, Erica, and Ulex. It has a preference for trunks and main stems ra<strong>the</strong>r than<br />
small twigs and branches. Amongst <strong>the</strong> <strong>British</strong> collections associated lichens on bark include<br />
Arthonia spadicea, Biatorina atropurpurea, 'Botrydina vulgaris', Cladonia spp., Hypogymnia<br />
physodes, Lecanora pallida, L. symmicta agg., Lecidea icmalea, Leparia incana agg., Micarea<br />
alabastrites, Micarea cinerea, Normandina pulchella, Ochrolechia androgyna, Farmelia saxatilis,<br />
Parmeliella jamesii, Pertusaria hymenea, Phyllopsora rosei, Platismatia glauca, Stenocybe<br />
septata and Trapelia sp. In <strong>the</strong> north and west <strong>of</strong> Britain it is frequently encountered on <strong>the</strong><br />
lignum <strong>of</strong> fallen trunks and large branches (especially conifers), accompanied by such species as<br />
Lecidea granulosa agg. , L. turgidula, Micarea denigrata, M. lignaria (including var. endoleuca),<br />
Mycoblastus sterilis, Ptychographa xylographoides, Xylographa abietina, and X. vitiligo. In <strong>the</strong><br />
same areas it is <strong>of</strong>ten common on peaty soil, and on moribund bryophytes or peaty debris on old<br />
walls, boulders and rock faces, preferring ra<strong>the</strong>r more sheltered conditions than M. lignaria,<br />
with which it is easily confused in <strong>the</strong> field. Fur<strong>the</strong>rmore, it is not found at such high altitudes as<br />
attained by that species, and all <strong>British</strong> collections seem to have been made at below 500 m.<br />
Associated species in <strong>the</strong>se habitats include Cladonia crispata, C. coccifera, C. squamosa, C.<br />
uncialis, Coelocaulon aculeatum s. lat., Hypogymnia physodes, Lecidea icmalea, Micarea<br />
leprosula, Ochrolechia androgyna, Farmelia saxatilis, dind Platismatia glauca. It is less frequent-<br />
ly found growing directly on rock, and records from Britain indicate that it is able to do so only in<br />
ra<strong>the</strong>r dry situations. Such occurrences are mostly in <strong>the</strong> ra<strong>the</strong>r low rainfall districts <strong>of</strong>, for<br />
example, Durham, east Yorkshire and Sussex; accompanying species noted include Baeomyces<br />
rufus, Cystocoleus ebeneus, Lecidea granulosa agg. , L. icmalea, Lecanora polytropa, Lepraria<br />
incana agg., Farmeliopsis ambigua, Trapelia coarctata, and T. involuta.<br />
M. peliocarpa is widely distributed in Britain but is most common in <strong>the</strong> west and in upland<br />
districts (but at low altitudes). This pattern is reflected in Europe as a whole, and it is apparently<br />
common in countries along <strong>the</strong> Atlantic seaboard and on <strong>the</strong> Atlantic islands including Iceland,<br />
<strong>the</strong> Azores, and Canary Islands. In central and eastern Europe it is found mainly in mountainous<br />
districts. Northwards it extends to just beyond <strong>the</strong> Arctic Circle, but I do not know <strong>of</strong> its<br />
occurrence in <strong>the</strong> high arctic. From outside Europe I have seen material from eastern Canada<br />
and north-eastern USA, and from New Zealand.<br />
Exsiccata: Anzi Lick. Sondr. 170A (UPS). Arnold Lich. Exs. 167A, B (BM ex K, M), 837, 1051 (BM ex<br />
K). Arnold Lich. Mon. 1 18 (BM ex K), 269, 357, 482 (BM ex K, MANCH). Hepp Flecht. Eur. 284 (E, M),<br />
285 (E, L, M), 510 (E, M). Hepp Zur. 206 (BERN). Korber Lich. Sel.Germ. 133A, B (M). Krypt. Exs.<br />
Vindob. 165 (BM). Larb. Lich. Herb. 347 (BM). Leighton Lich. Brit. 23^ p.p. (BM, DBN, FRS). Lojka<br />
Lich. Hung. 134 (BM ex K). Malme Lich. Suec. 169 (S). Rasanen Lichenoth. Fenn. 343 (BM). Vezda Lich.<br />
Sel. 1342, 1380 (BM). Zwackh. Lich. Exs. 276, 897 (UPS).
34. Micarea prasina Fr.<br />
LICHEN GENUS MICAREA IN EUROPE 173<br />
-^^r<br />
Map 17 Micarea peliocarpa # 1950 onwards O Before 1950<br />
(Figs ID, 26-27, 49, 53-54; Maps 18-20)<br />
Syst. orb. : 256-7 (Dec. 1825). - Biatora prasina Fr. , Stirp. agrifemsion.: 36 (June 1825), nom. illeg. (Art.<br />
63). - Biatora prasina (Fr.) Trevisan, Linnaea 28: 288 (1856), nee Tuck. & Mont., in Mont. (1857). -<br />
Catillaria prasina (Fr.) Th. Fr., Lich. Scand. 2: 572 (1874). See note (i) below and note on Sporacestra<br />
under 'Excluded taxa'.<br />
Biatora micrococca Korber, Parerg. lich.: 155 (1860). - Catillaria micrococca (Korber) Th. Fr., Lich.<br />
Scand. 2: 571 (1874). - Micarea prasina f. micrococca (Korber) Hedl. in Bih. K. svenska VetenskAkad.<br />
Handl. Ill, 18 (3): 77, 87 (1892). - Micarea micrococca (Korber) H. Gams, Kleine Kryptfl. 3: 67 (1967);<br />
comb, inval. (Art. 33.2). Type: Germany, Baden-Wiirttemberg, 'in regno Wurtembergico', on Pinus<br />
bark, 1862, K. A. Kemmler, Korber Lich. Sel. germ. 250 (L 910, 139-1361-neotype! [t.l.c: 'prasina-<br />
unknown A']). See note (ii) below.<br />
Lecidea subviridescens Nyl. in Flora, Jena 51: 474 (1868) . - Micarea subviridescens (Nyl.) Hedl. in Bih. K.<br />
svenska VetenskAkad. Handl. Ill, 18 (3): 77, 87 (1892). - Bacidia subviridescens (Nyl.) Zahlbr., Cat.<br />
lich. univ. 4: 154 (1926). Type: Channel Islands, Jersey, Boulay Bay, on coastal turf and soil, 1868, C.<br />
Larbalestier (H-NYL 19056 [as 'subvirescens'] - lectotype! ; BM - isolectotype! [t.l.c. : 'prasina unknown<br />
C']; BM-topotypes!). See note (iii) below.<br />
Lecidea prasiniza Nyl. in Flora, Jena 57: 312 (1874). - Micarea prasina f. byssacea subf . prasiniza (Nyl.) Th.<br />
Fr. in Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 77 (1892). Type: Finland, Tavastia<br />
australis, Padasjoki, Nyystola, on Alnus, 1872, E. A. Lang fVainio] (H-NYL 21604 - lectotype!, H -<br />
isolectotype!). [material insufficient for t.l.c.].<br />
Lecidea sordidescens Nyl. in Flora, Jena 57: 312 (1874). - Lecidea erysiboides f. sordidescens Nyl. in<br />
Norrlin in Not. Sdllsk. Fauna Fl. fenn. Forh. 11: 188 (1870); nom nudum (Art. 32). - Catillaria prasina
174 BRIAN JOHN COPPINS<br />
var. byssacea f . sordidescens (Nyl.) Blomb. & Forss. , Enum. PI. Scand. : 91 (1880). - Micarea prasina f<br />
byssacea subf. sordidescens (Nyl.) Th. Fr. in Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3):<br />
77 (1892). - Lecidea byssacea var. sordidescens (Nyl.) Vainio in Termeszetr. Fuz. 22: 320 (1899). -<br />
Catillaria prasina var. sordidescens (Nyl.) Lettau in Hedwigia 52: 136 (1912). - Micarea prasina var.<br />
sordidescens (Nyl.) Brodo in Bull. N. Y. St. Mus. Sci. Serv. 410: 152 (1968). Type: Switzerland, Zurich,<br />
'An faulen Fichten-Strunken Z. H.', Hepp, Flecht. Eur. 11^ (E - lectotype! [t.l.c.: 'prasina-unknown<br />
B']; isolectotypes: BM (boxed set)! [t.l.c: 'prasina-unknown B'], H-NYL 21632 [fragment]!, M!). See<br />
note (iv) below.<br />
Lecidea prasiniza var. prasinoleuca Nyl. in Flora, Jena 64: 7 (1881). Type: Germany, Baden-Wiirttemberg,<br />
Heidelberg, Konigstuhle, on Picea abies, ix 1880, Zwackh, Lich. Exs. 593a (H-NYL 21601 - lectotype!<br />
[t.l.c: 'prasina-unknown A']). See note (v) below.<br />
Catillaria micrococca f. glebosula Erichsen in Annls mycol. 36: 139 (1938). Type: Germany, Schleswig-<br />
Holstein, Eiderstedt, Ording, at base <strong>of</strong> Betula, 4 viii 1913, C. F. E. Erichsen (HBG - holotype! [t.l.c:<br />
'prasina-unknown B'].<br />
Catillaria micrococca var. discrepans Erichsen in Annls mycol. 36: 139 (1938). Type: Denmark, Jylland,<br />
Abenra (Apenrade), on smooth bark oiAlnus in a valley near Elisenlund, 3 ix 1913, C. F. E. Erichsen<br />
(HBG - holotype!). [material insufficient for t.l.c].<br />
Catillaria prasina var. occulta Erichsen in Annls mycol. 36: 140 (1938). Type: Germany, Schleswig-<br />
Holstein, Ratzeburg, in <strong>the</strong> wood 'Bak', on ± smooth bark oiAlnus, 21 iii 1918, C. F. E. Erichsen (HBG<br />
- holotype!). [material insufficient for t.l.c].<br />
Lecidea declivitatum Erichsen in Mitt. Inst. allg. Bot. Hamb. 10: 408 (1939). Type: Germany, Schleswig-<br />
Holstein, 'Krs Eutin: an der Boschung eines Waldwegs an der Nordwest-Seite des Ugleisees', 5 vi 1914,<br />
C. F. E. Erichsen (HBG -holotype! [t.l.c: 'prasina-unknown C']).<br />
Micarea polytrichi Poelt & Dobb. in Bot. Jb. 96: 343 (1975). Type: Austria, Steiermark, Grazer Bergland,<br />
a little north <strong>of</strong> Maria Trost near <strong>the</strong> Wenisbucher Strasse, c. 460 m, on Polytrichumformosum on shady<br />
side <strong>of</strong> a narrow gorge, 30 iii 1974, /. Poelt (GZU - holotype!) [material insufficient for t.l.c].<br />
Catillaria prasina a laeta Th. Fr., Lich. Scand. 2: 573 (1874); nom. inval. (Art. 26).<br />
?Biatora byssacea Zwackh in Flora, Jena 45: 510 (1862), non Hampe in Linnaea 25: 709 (1852). Type:<br />
Germany, Heidelberg, 'Konigsstuhle', on bark <strong>of</strong> old Quercus, ? Zwackh (not traced). See note (vi)<br />
below.<br />
?Catillaria melanobola f. biseptata B. de Lesd., Rech. Lich. Dunkerque: 198 (1910). Type: France, Nord,<br />
Dunkerque, St Pol, on a piece <strong>of</strong> lea<strong>the</strong>r in dunes, B. de Lesdain (not seen).<br />
?Catillaria melanobola f. nigra B. de Lesd., Rech. Lich. Dunkerque: 199 (1910). Type: France, Nord,<br />
Dunkerque, lignum <strong>of</strong> hollow Salix, B. de Lesdain (not seen).<br />
Notes: (i) Biatora prasina Fr. (June 1825) is superfluous because <strong>the</strong> protologue includes <strong>the</strong> sentence:<br />
'Crusta est Byss. botryoides Linn!'. Byssus botryoides L. is a valid name {Sp. pi. 2: 1169 (1753)) but <strong>the</strong><br />
material in <strong>the</strong> Linnaean herbarium (LINN) is a specimen (1278.16) labelled by Ehrhart; it is an alga and<br />
bears a determination label: 'Protococcus viridis Ag. Determined by Francis Drouet iv. 1950'. In <strong>the</strong><br />
protologue <strong>of</strong> Micarea and M. prasina. Fries (Dec. 1825) makes a less dogmatic statement, merely<br />
indicating that when sterile M. prasina is indistinguishable from B. botryoides; M. prasina is, <strong>the</strong>reby, not<br />
rendered superfluous.<br />
(ii) I have seen two specimens named by Korber and collected from 'Wiirtemberg' by Kemmler: an<br />
undated specimen in WRSL and possibly part <strong>of</strong> <strong>the</strong> original ga<strong>the</strong>ring, and a later specimen issued in<br />
Korber's exsiccata. The first specimen would have to be completely destroyed for t.l.c. analysis, whereas<br />
<strong>the</strong> exsiccate specimen is larger and was found to contain 'prasina-unknown A', and it is this specimen that I<br />
have chosen for typification. If future studies require this chemical race to be recognized at specific rank,<br />
<strong>the</strong>n 'micrococca' is <strong>the</strong> earliest available epi<strong>the</strong>t.<br />
(iii) L. subviridescens is <strong>the</strong> earliest name for <strong>the</strong> chemical race with 'prasina-unknown C.<br />
(iv) Although Nylander (1874) did not provide a description, he validated L. sordidescens by referring to<br />
it Hepp Flecht. Eur. 278; <strong>the</strong> printed label for this exsiccatum includes a short description and a drawing <strong>of</strong><br />
spores.<br />
(v) If <strong>the</strong> chemical race containing 'prasina-unknown A' were to be awarded varietal rank, <strong>the</strong>n this<br />
name provides <strong>the</strong> earliest available epi<strong>the</strong>t, provided that 'byssacea' is not available; see also note (vi).<br />
(vi) I have not seen a holotype specimen or any material suitable for <strong>the</strong> lectotypification <strong>of</strong> Biatora<br />
byssacea Zwackh. A neotype has not been selected because this name is a later homonym <strong>of</strong> B. byssacea<br />
Hampe. Hampe's name was given to a specimen collected by Charles Stuart in Tasmania, and <strong>the</strong><br />
protologue ends with 'an status degenerans'. Biatora byssacea Hampe is included by Zahlbruckner (Cat.<br />
lich. univ. 3: 897) in a list <strong>of</strong> excluded and dubious names and is not accorded a catalogue number; <strong>the</strong> type<br />
.
LICHEN GENUS MICAREA IN EUROPE 175<br />
material has not been located in BM but it could be elsewhere (? G). Under <strong>the</strong> taxonomic arrangement<br />
given here Zwackh's epi<strong>the</strong>t is <strong>of</strong> no consequence. If <strong>the</strong> chemical races <strong>of</strong> M. prasina were to be awarded<br />
varietal status, <strong>the</strong>n under <strong>the</strong> provisions <strong>of</strong> Article 72 <strong>the</strong> name ' Catillaria prasina p. [var.] byssacea Th. Fr.<br />
(1874)' could become important and require typification. If this name was typified with a specimen<br />
containing 'prasina unknown A' <strong>the</strong>n it would be <strong>the</strong> earliest available name at that rank.<br />
Thallus effuse and <strong>of</strong>ten wide-spreading, very variable in colour, pale fawn (dry shaded<br />
situations), light green, pale to dark grey-green (<strong>of</strong>ten with glaucous hue), or olivaceous to<br />
olive-black, <strong>of</strong>ten appearing ± gelatinous when moist, composed <strong>of</strong> small ± globose granules<br />
(goniocysts). Goniocysts c. 12^0(-60) jxm diam, thinly scattered, or vertically proliferating to<br />
give <strong>the</strong> thallus a s<strong>of</strong>t isidiose appearance, or densely aggregated to form a thick granular crust<br />
up to 200 />tm deep. Goniocysts ecorticate, <strong>of</strong>ten with shortly protruding hyphal filaments;<br />
outermost hyphae <strong>of</strong> superficial goniocysts sometimes surrounded by greenish, K+ violet<br />
pigment. Phycobiont micareoid, cells 4-7 ^tm diam.<br />
Apo<strong>the</strong>cia usually numerous but sometimes few or absent, immarginate, sometimes adnate<br />
and shallow-convex, but more usually becoming markedly convex, subglobose or tuberculate;<br />
colour variable, whitish or pallid {'micrococca' shade-forms), pale to dark grey, brownish grey,<br />
or grey-black, matt; 0- 1-0-4 mm diam, or to 0-6 mm when tuberculate. Hymenium 28-45(-52)<br />
H^va tall, hyaline or dilute dull straw, but more usually dull olivaceous, especially in <strong>the</strong> upper<br />
part (but coloured portion not sharply delimited) or in vertical streaks; pigment K+ violet,<br />
HNO3+ red, confined to gel-matrix. Asci clavate or cylindrical-clavate, 26-40(-50)x8-12 /u,m.<br />
Spores ovoid-ellipsoid, ovoid, ovoid-oblong or oblong, 0-l(-3)-septate; when 1-septate <strong>the</strong><br />
lower cell is slightly longer and narrower than <strong>the</strong> upper, and <strong>the</strong>re is <strong>of</strong>ten a slight constriction at<br />
<strong>the</strong> septum; variable in size, in range 8-14(-17)x2-3-4(-5) /xm. Paraphyses ra<strong>the</strong>r numerous,<br />
branched and anastomozing, 0-5-1 /xm wide; apical parts <strong>of</strong>ten wider, to 1-5 /xm, but not<br />
thickened by pigment; epi<strong>the</strong>cial pigment confined to surrounding gel matrix. Hypo<strong>the</strong>cium c.<br />
40-170 /xm tall, hyaline to dilute dull yellowish; hyphae interwoven, some becoming vertically<br />
orientated towards <strong>the</strong> hymenium, c. 0-7-1-3 jxm wide; ascogenous hyphae with swollen cells, c.<br />
2-4 /xm wide. Excipulum poorly developed, sometimes evident as a narrow, hyaline or dilute<br />
olivaceous-straw (K-l- violet), non-amyloid, reflexed zone; hyphae radiating, branched and<br />
anastomosing, c. 0-7-1 /am wide.<br />
Pycnidia <strong>of</strong>ten present but usually inconspicuous, white, or upper part around <strong>the</strong> ostiole grey<br />
(due to olivaceous, K+ violet pigment), immersed to sessile (never distinctly stalked), <strong>of</strong> two<br />
types: (a) c. 50-120 /xm diam, emergent to sessile, ostiole sometimes gaping; conidia (mesoconi-<br />
dia) ± cylindrical or narrowly obpyriform, <strong>of</strong>ten biguttulate and slightly constricted near <strong>the</strong><br />
middle, (3-5-)4-6x 1-2-1-7 /xm; (b) c. 30-60(-100) /xm, usually immersed in surrounding<br />
goniocysts, ostiole rarely gaping; conidia (microconidia) cylindrical or narrowly fusiform,<br />
(5-)5-5-8x0-7-l/xm.<br />
Chemistry: All parts K— , C—<br />
(but olivaceous pigment C-l- violet, never red), PD— ; t.l.c:<br />
specimens have one <strong>of</strong> three unknowns ('prasina unknown A', 'B', or 'C; see p. 87); trace<br />
amounts <strong>of</strong> gyrophoric acid sometimes detected, possibly due to contamination by intimately<br />
associated species such as Lecidea icmalea and L. granulosa agg.<br />
Observations: Micarea prasina is <strong>the</strong> commonest and most variable member <strong>of</strong> <strong>the</strong> genus.<br />
However, it is one <strong>of</strong> <strong>the</strong> few European species whose thallus is composed <strong>of</strong> minute discrete<br />
granules (goniocysts). Two o<strong>the</strong>r species with a similar thallus are: M. hedlundii (q.v.) with a<br />
K-l- red oily substance in <strong>the</strong> goniocysts and conspicuous tomentose, brown, stalked pycnidia;<br />
and M. melanobola which has a sharply delimited epi<strong>the</strong>cium (pigment closely adhering to<br />
apices <strong>of</strong> <strong>the</strong> paraphyses) and shorter microconidia . Both <strong>of</strong> <strong>the</strong>se species appear to be very rare<br />
A comparison <strong>of</strong> <strong>the</strong> diagnostic features <strong>of</strong> M. melanobola, M. prasina and M. misella are given<br />
in Table 6.<br />
Diminutive forms <strong>of</strong> M. prasina with small pallid apo<strong>the</strong>cia and a scanty thallus have<br />
sometimes been regarded as a distinct species {'Catillaria micrococca') .<br />
Such forms have <strong>the</strong><br />
same basic thallus structure, spores, paraphyses, anamorphs and chemistry (with ei<strong>the</strong>r 'prasina<br />
unknown A' or 'B') as more typical forms (with coloured apo<strong>the</strong>cia and more robust thallus).<br />
.
176 BRIAN JOHN COPPINS<br />
and I have seen intermediates on many occasions. The 'micrococca' forms are found in heavily<br />
shaded situations (e.g. tree trunks in dense conifer plantations, and trunks <strong>of</strong> understory trees or<br />
shrubs) and I consider <strong>the</strong>ir diminutive stature and lack <strong>of</strong> pigmentation to be a phenotypic<br />
response to low light intensities. The specimens described as Micarea polytrichi are similarly<br />
diminished forms, in this case overgrowing mosses (mainly Polytrichum spp.) by woodland<br />
roads.<br />
European populations <strong>of</strong> M. prasina have been found to be represented by three chemical<br />
races, each with one <strong>of</strong> three distinctive, but as yet unidentified, substances (see p. 87). During<br />
<strong>the</strong> course <strong>of</strong> <strong>the</strong> present study I have been tempted to recognize <strong>the</strong> chemical races as formal<br />
varieties. However I defer from doing so at this time because: (a) <strong>the</strong> chemical structure and<br />
biogenetic relationships between <strong>the</strong> three substances is not yet known; (b) <strong>the</strong> three chemical<br />
races do not consistently correlate to any clear differences in morphology; (c) <strong>the</strong> correlations<br />
between chemistry and distribution and (or) ecology are as yet uncertain. In <strong>the</strong> <strong>British</strong> Isles (at<br />
least) <strong>the</strong> race containing <strong>the</strong> 'unknown A' is by far <strong>the</strong> commonest and is a ubiquitous coloniser<br />
<strong>of</strong> trees and shrubs in all types <strong>of</strong> woodland (including young conifer plantations), occurring on<br />
tree boles, bark and lignum <strong>of</strong> fallen trees and stumps, and fallen sticks; it also occurs on shaded<br />
mossy turf on <strong>the</strong> ground and amongst rocks (usually with an east- or north-facing aspect) in<br />
maritime situations. The race containing 'unknown B' has a more restricted ecology and is<br />
mainly found in old woodland situations (including <strong>the</strong> native pine-woods), growing on <strong>the</strong><br />
ra<strong>the</strong>r dry lignum or old bark <strong>of</strong> large stumps or fallen trunks; it has not been found on coastal<br />
turf or rocks. Specimens <strong>of</strong> this race tend to have a thick, pale grey (<strong>of</strong>ten abrading to greenish<br />
Map 18 Micarea prasina sensu lato • 1950 onwards O Before 1950
LICHEN GENUS MICAREA IN EUROPE 177<br />
Map 19 Micarea prasina substance A # 1950 onwards O Before 1950<br />
white) thallus, but equivalent morphological forms containing 'unknown A' are sometimes<br />
encountered. Specimens containing 'unknown C are exceedingly rare and are known only from<br />
coastal turf in south-west England and <strong>the</strong> Channel Islands, and on argillaceous soil on a bank by<br />
a woodland path in south-eastern Schleswig-Holstein. The above ecological and phytogeographical<br />
tendencies require to be more thoroughly tested throughout <strong>the</strong> range <strong>of</strong> M. prasina,<br />
and it is possible that a good case for <strong>the</strong> taxonomic recognition <strong>of</strong> <strong>the</strong>se races as varieties,<br />
subspecies, or even species could be made.<br />
The type material <strong>of</strong> Lecidea subviridescens contains 'unknown C and has spores that are at<br />
<strong>the</strong> high end <strong>of</strong> <strong>the</strong> overall size range for M. prasina s. ampl. (10-18x4-5-5 /xm) and some with<br />
two or three septa. However <strong>the</strong> very large 2-3-septate spores are always old with secondarily<br />
thickened walls, and similar spores are sometimes encountered in o<strong>the</strong>r Micarea species that<br />
have predominantly 1-septate spores. Fur<strong>the</strong>rmore, similarly large, 2-3-septate spores have<br />
been found in some coastal specimens containing 'unknown A' and in some inland, lignicolous<br />
specimens containing 'unknown B'. Although I regarded 'subviridescens'' as a separate species in<br />
<strong>the</strong> recent <strong>British</strong> checklist (Hawksworth etal., 1980: 62), I now believe it should be subsumed<br />
under M. prasina pending fur<strong>the</strong>r studies as intimated in <strong>the</strong> preceding paragraph.<br />
Habitat and distribution (see also 'observations'): As an epiphyte on trees or shrubs M.<br />
prasina has been found on a wide range <strong>of</strong> phorophytes; in Britain <strong>the</strong>se include Acer, Alnus,<br />
Betula, Castanea, Corylus, Fagus, Fraxinus, Metrosideros, Quercus, Salix, Sambucus, Sorbus,<br />
Ulmus, Calluna, Erica, Rhododendron, Vaccinium, Sarothamnus, Ulex, Abies, Larix, Picea^<br />
Pinus, Pseudotsuga, and Juniperus. It occurs on <strong>the</strong> bark or lignum (old dried wounds, etc.) <strong>of</strong>
178 BRIAN JOHN COPPINS<br />
Map 20 Micarea prasina substance B # 1950 onwards O Before 1950 substance C ® Before 1950 I<br />
shaded trunks or main stems, but also on low (shaded) branches, fallen debris (especially in<br />
conifer plantations) and stumps. It is usually found in woodlands but sometimes occurs in<br />
sheltered niches in situations, e.g. bases <strong>of</strong> Ulex stems in more open, exposed hillside<br />
gorse-scrub.<br />
The phytosociological affinities <strong>of</strong> M. prasina are various and difficult to define. When on <strong>the</strong><br />
bark <strong>of</strong> deciduous trees it sometimes occurs in shaded facies <strong>of</strong> communities belonging to <strong>the</strong><br />
Lobarion pulmonariae, Parmelietum revolutae, and <strong>the</strong> Pseudevernietum furfuraceae. It is<br />
frequently found in small bark crevices amongst ± smooth areas <strong>of</strong> bark which are colonized by<br />
communities <strong>of</strong> <strong>the</strong> Graphidetum scriptae, Pyrenuletum nitidae, and Lecanoretum subfuscae.<br />
When occurring in such situations as <strong>the</strong> shaded trunks <strong>of</strong> young conifers or shrubs M. prasina<br />
<strong>of</strong>ten exists as ± pure stands; more detailed phytosociological studies may well lead to <strong>the</strong><br />
formal syntaxonomic recognition <strong>of</strong> such communities. From amongst <strong>the</strong> <strong>British</strong> collections in<br />
E <strong>the</strong> associated species <strong>of</strong> M. prasina on <strong>the</strong> bark <strong>of</strong> deciduous trees include Arthonia spadicea,<br />
Bacidiavezdae, Chrysothrix candelaris, Cladonia coniocraea, Dimerella diluta, Evernia prunas-<br />
tri, Graphis elegans, Gyalideopsis anastomosans, Hypogymnia physodes, Lecidea icmalea,<br />
Lepraria incana, Ochrolechia androgyna, Opegrapha vulgata, Pachyphiale cornea, Parmelia<br />
glabratula, P. sulcata, Pertusaria multipuncta, Thelotrema lepadinum, Trapelia corticola, Dicranoweissia<br />
cirrata, Frullania spp., Hypnum cupressiforme, Lejeunea ulicina, and Metzgeria<br />
fareata.<br />
When on lignum its affinities are similarly difficult to define but it is <strong>of</strong>ten found in more<br />
sheltered or shaded (humid) niches adjacent to communities <strong>of</strong> <strong>the</strong> Calicietum abietini,<br />
Cladonietum coniocraeae, Lecideetum ostreatae, and Parmeliopsidetum ambiguae.
LICHEN GENUS MICAREA IN EUROPE 179<br />
Species associated with M. prasina on <strong>the</strong> hgnum <strong>of</strong> fallen trunks and large stumps in <strong>the</strong><br />
<strong>British</strong> Isles include Bryoria fuscescens, Chaeno<strong>the</strong>ca ferruginea, Chaeno<strong>the</strong>copsis spp., Cladonia<br />
chlorophaea agg., C. macilenta, C. ochrochlora, Hypocenomyce scalaris, Hypogymnia<br />
physodes, Lecanactis abietina, Lecidea aeruginosa, L. icmalea, Micarea adnata, M. melaena,<br />
and Platismatia glauca.<br />
In coastal districts M. prasina is <strong>of</strong>ten found on plant debris, soil or moribund bryophytes in<br />
rock crevices or on ledges in sheltered gullies, and sometimes it occurs on <strong>the</strong> ground growing<br />
over plant debris or old Armeria tussocks, etc. Such habitats are usually sheltered and (or) with a<br />
north- to east-facing aspect. Associated lichens are few, but include Cladonia spp., Lecidea<br />
granulosa, L. icmalea, and Lepraria incana agg. Finds <strong>of</strong>M. prasina growing directly on rock are<br />
very rare, but I have seen a few collections made from sandstone rocks in woodlands (from east<br />
Sussex and north-east Yorkshire).<br />
M. prasina is still to be found close to <strong>the</strong> centres <strong>of</strong> large conurbations and cities (e.g. Bristol,<br />
Edinburgh, London, and <strong>the</strong> West Yorkshire conurbation), and its persistence is probably due<br />
more to its ability to avoid, ra<strong>the</strong>r than tolerate, <strong>the</strong> direct effects <strong>of</strong> air pollutants such as<br />
sulphur dioxide and its derivatives. It naturally favours substrata with a low pH and is able to<br />
grow in very sheltered and shaded situations where more light demanding pollution resistant<br />
species (especially Lecanora conizaeoides) are at a competitive disadvantage.<br />
M. prasina s. ampl. is widely distributed in <strong>the</strong> <strong>British</strong> Isles and much <strong>of</strong> Europe. In<br />
Scandinavia it occurs northwards to at least c. 67°N. It may be rare in districts adjoining <strong>the</strong><br />
Mediterranean Sea, but those areas are mostly poorly known lichenologically, and I have seen<br />
one collection from Toscana (nor<strong>the</strong>rn Italy). M. prasina is present in Macaronesia (Azores and<br />
Canary Islands) and I have seen collections <strong>of</strong> it from several states in <strong>the</strong> U.S.A. (Georgia,<br />
Maryland, Massachusetts, Michigan, and Wisconsin). In addition I have seen several collections<br />
from South America and Australisia which come close to M. prasina, but <strong>the</strong>se require more<br />
critical study and will be treated in a later publication.<br />
Exsiccata. Containing 'prasina unknown A': Arnold Lich. Exs. 1122, 1472 (BM ex K, M). Korber Lich<br />
Sel. Germ. 250 (L). Lojka Lich. Univ. 29 (BM ex K, M), 30 (BM ex K). Malme Lich. Suec. 23 (M, S)<br />
Rabenh. Lich. Eur. 676 (BM, BM ex K, M, WRSL). Rasanen Lich. Fenn. 651 (BM, BM ex K, M), 652<br />
(BM, BM ex K). Vezda Lich. Sel. 90 (BM, M), 1467 (BM). Zwackh. Lich. Exs. 593A (H-NYL 21601)<br />
Containing 'prasina unknown B': Arnold Lich. Exs. 280C (BM, GZU, M, WRSL). Cumm. Dec. N. Am<br />
Lich. I. 355 (BM, WIS). Hepp Flecht. Eur. 278 (BM, E, M, WRSL). Lojka Lich. Univ. 31 (BM ex K, M)<br />
Chemistry not tested: Arnold Lich. Exs. 279 (WRSL), 280A (BM, BM ex K, M), 280B (BM ex K, M)<br />
Arnold Lich. Mon. 243 (UPS), 245 (BM ex K, M). Hepp Zur. 224 (BERN). Kutak Lich. Bohem. 310 (O)<br />
Magnusson Lich. Sel. Scand. 134 (BM). Malme Lich. Suec. 24 (M, S). Rabenh. Lich. Eur. 733 (H)<br />
Rasanen Lich. Fenn. 653 (LD). Schaerer Lich. Helv. 196 p.p. (BM ex K, M). Vezda Lich. Sel. 1595 (BM<br />
GZU, M). Zahlbr. Lich. Rar. 175 (BM). Zwackh. Lich. Exs. 416 (UPS), 591A (H-NYL 21598), 591B<br />
(H-NYL 21594, 21599), 592A (UPS), 592B (M, UPS), 592C (UPS), 592D (M, UPS), 592E (UPS), 593B<br />
(H-NYL 21600), 593C (H-NYL 21595), 656 (UPS).<br />
35. Micarea pycnidiophora Coppins & P. James<br />
(Fig. 28: Map 21)<br />
in Lichenologist 11: 153 (1979). Type: England, South Hampshire, New Forest, near Cadnam, Shave<br />
Wood, 45 m, on bark <strong>of</strong> Fagus, 5 xi 1972, B. J. Coppins & F. Rose (E - holotype!; BM - isotype!).<br />
Thallus corticolous, <strong>of</strong>ten overgrowing moribund thaUi <strong>of</strong> bryophytes and o<strong>the</strong>r hchens;<br />
effuse, <strong>of</strong>ten wide-spreading, thin, uneven, composed <strong>of</strong> scattered to confluent areolae arising<br />
from a thin varnish-like prothallus. Areolae flattened to convex-hemispherical, grey-green or<br />
dull green, c. 40-100 ^im diam; in section, without a distinct cortex or hyaline amorphous<br />
covering layer. Phycobiont micareoid, cells 4-7 /^m diam.<br />
Apo<strong>the</strong>cia usually few or absent but sometimes abundant, immarginate, convexhemispherical<br />
to globose, <strong>of</strong>ten tuberculate, ivory-white to palUd, translucent when moist,<br />
surface matt, 0- 1-0-3 mm diam. Hymenium 35-50 /xm, hyaUne. Asci clavate, 30-35 x 10-12 /am.<br />
Spores ± straight and vertically aligned in <strong>the</strong> ascus (never tightly spiralled), shortly acicular.
180 BRIAN JOHN COPPINS<br />
Upper end usually broader and more obtuse than <strong>the</strong> lower end, usually slightly curved,<br />
3-7-septate, (14-)21-34x2-2-5(-2-7) (xm. Paraphyses numerous, branched and anastomosing,<br />
c. 1-1-5 ^tm, not swollen at apices. Hypo<strong>the</strong>cium 20-80 ^tm tall, hyaline; hyphae interwoven, but<br />
becoming vertically orientated towards <strong>the</strong> hymenium, c. 1-1-5 /xm wide; ascogenous hyphae<br />
with swollen cells, c. 2-4 /xm wide. Excipulum poorly developed (c. 10 /am wide); hyphae<br />
radiating, branched and anastomosing, c. 1 /xm wide.<br />
Pycnidia always numerous and conspicuous, sessile or shortly stalked, whitish (concolorous<br />
with <strong>the</strong> apo<strong>the</strong>cia), 100-300 /xm tall (including stalk) and 60-120 /xm diam; stalks (pycnidiophores)<br />
simple but occasionally clustered and appearing as if branched at <strong>the</strong> base; stalk<br />
tissue composed <strong>of</strong> interwoven hyphae c. 1-1-5 /am wide that ± separate in K. Conidiogenous<br />
cells ± cylindrical, 5-10x1-1-5 (xm. Conidia (mesoconidia) cylindrical, eguttulate, 3-8-6x1l-2(-l-5)/xm.<br />
acid.<br />
Chemistry: Apo<strong>the</strong>cia, pycnidia and thallus (in section) K-, C+ red, PD-; t.l.c: gyrophoric<br />
Observations: Micarea pycnidiophora is characterized by its whitish apo<strong>the</strong>cia and concol-<br />
orous, ± stalked pycnidia, shortly acicular spores, and C-l- red reactions (gyrophoric acid). It<br />
most closely resembles M. stipitata, which differs in having more elongate and <strong>of</strong>ten distinctly<br />
branched pycnidiophores, larger conidia, and C— reactions (gyrophoric acid absent). M.<br />
globulosella and M. syno<strong>the</strong>oides have similar acicular spores, but <strong>the</strong>ir apo<strong>the</strong>cia are darkcoloured<br />
with an olivaceous (K-l- violet) pigment, and <strong>the</strong>ir pycnidia are inconspicuous and<br />
usually immersed in <strong>the</strong> thallus. Confusion could arise with Scoliciosporum pruinosum (P.<br />
Map 21 Micarea pycnidiophora k. + Micarea stipitata^
LICHEN GENUS MICAREA IN EUROPE 181<br />
James) Vezda and 5". schadeanum (Erichsen) Vezda, but those species have narrower spores<br />
which are spiralled in <strong>the</strong> ascus and non-micareoid phycobionts (? Trebouxia); moreover, S.<br />
pruinosum has a granular epi<strong>the</strong>cium (granules dissolving in K). Fur<strong>the</strong>rmore, <strong>the</strong>se species<br />
have inconspicuous (? unknown) pycnidia, and do not contain gyrophoric acid.<br />
For illustrations <strong>of</strong> conidia and conidiogenous cells, and for fur<strong>the</strong>r discussions, see Coppins<br />
& James (1979).<br />
Habitat and distribution: M. pycnidiophora is a species <strong>of</strong> old woodlands, in damp, sheltered,<br />
and ra<strong>the</strong>r shaded situations. It is generally found on areas <strong>of</strong> ± smooth-bark on <strong>the</strong> boles <strong>of</strong> old<br />
trees, especially Fagus and Ilex, but also Alnus, Quercus, and Rhododendron. Although<br />
characteristic <strong>of</strong> acid bark it has not yet been recorded on conifers.<br />
Its known centre <strong>of</strong> distribution is <strong>the</strong> New Forest <strong>of</strong> Hampshire in sou<strong>the</strong>rn England, where it<br />
is locally frequent. Its o<strong>the</strong>r <strong>British</strong> localities are Sheffield Park in Sussex (on Rhododendron<br />
with Gyalideopsis anastomosans) and <strong>the</strong> Limb Valley, Yorkshire (its most nor<strong>the</strong>rly locality; at<br />
base <strong>of</strong> Fraxinus). The latter site is an ancient oak-wood just outside <strong>the</strong> large industrial city <strong>of</strong><br />
Sheffield, but M. pycnidiophora occurs in an extremely sheltered situation, where it presumably<br />
avoids <strong>the</strong> effects <strong>of</strong> <strong>the</strong> high levels <strong>of</strong> air pollution prevalent in that area. M. pycnidiophora is<br />
little known outside <strong>of</strong> Britain, but is reported from Bretagne, France (Coppins, 1971, as<br />
'Bacidia sp. 2') and <strong>the</strong> Canary Islands.<br />
36. Micarea rhabdogena (Norman) Hedl.<br />
(Fig. 29A)<br />
in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 75, 85 (1892). - Biatora (Lecidea) rhabdogena<br />
Norman in Ofvers. K. VetenskAkad. Fork. 11: 803 (1870). - Lecidea rhabdogena (Norman) Th. Fr.,<br />
Lich. Scand. 2: 473 (1874). Type: Norway, Nordland, Maalselven, Skjeggenaes, J. M. Norman (O -<br />
lectotype!; isolectotypes: BM!, O!, S!).<br />
Lecidea glomerellaf. ecrustacea Nyl. ex Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 457 (1934). - Lecidea<br />
glomerella f. ecrustacea Nyl. in Elfving in Medd. Soc. Fauna Fl. fenn. 2: 168 (1878); nom. nudum (Art.<br />
32). Type: USSR, Karelskaya ASSR, Karelia olonetsensis, Gorki near Svir, F. Elfving (H-NYL 19120-<br />
holotype!).<br />
Thallus inapparent, endoxylic; hyphal walls <strong>of</strong>ten olivaceous, K+ violet. Phycobiont probably<br />
micareoid, cells c. 5-7 ixm diam.<br />
Apo<strong>the</strong>cia numerous, immarginate, convex-hemispherical or tuberculate, black, matt, 0-1-<br />
0-3 mm diam, or up to 0-4 mm diam when tuberculate. Hymenium 23-30 /xm tall; upper part<br />
(epi<strong>the</strong>cium) fuscous-brown, HNO3— , K- but pigment dissolving and fading into solution;<br />
remaining (lower) part dilute yellowish brown; hymenium sometimes tinged violet in K (evident<br />
after brown pigment has dissolved). Asci clavate, 19-28x7-9 /am. Spores oblong-ellipsoid,<br />
ovoid-oblong or bacilliform, simple or rarely a few 1-septate, 6-9xl-5-2-3 /am. Paraphyses<br />
numerous, hyaline throughout, c. 1 /xm wide in mid-hymenium but <strong>of</strong>ten widening above to T5<br />
/am; sparingly branched and sometimes anastomosing, but becoming richly branched above<br />
where <strong>the</strong>ir entangled apices, toge<strong>the</strong>r with <strong>the</strong> brown pigment, form a ± well delimited<br />
epi<strong>the</strong>cium. Hypo<strong>the</strong>cium c. 70-160 /am tall, pale, dilute yellowish brown, K— , HNO3—<br />
Excipulum reflexed, dilute to dark fuscous-brown, K— but partly dissolving into solution;<br />
hyphae radiating, branched and anastomosing, c. 0-7-OT /am wide, sometimes widening to 1-5<br />
/am towards <strong>the</strong> outer edge.<br />
Pycnidia <strong>of</strong> two types: (a) partly immersed to ± sessile, black, matt, 40-80 /am diam; wall<br />
olivaceous, K-l- violet; conidia (mesoconidia) oblong-ellipsoid, oblong-ovate or obovate, c.<br />
3-5-4-7X1-4—T8 /am; (b) immersed between surface wood fibres, black, c. 4 /am diam; wall<br />
olivaceous, K-l- violet; conidia (microconidia) narrowly cylindrical, 4-4-5 xO-7-0-9 /am.<br />
Chemistry: Sections <strong>of</strong> apo<strong>the</strong>cia C— ; material insufficient for analysis by t.l.c.<br />
Observations: M. rhabdogena resembles M. elachista in apo<strong>the</strong>cial pigmentation and anatomy,<br />
but differs in its endoxylic thallus, smaller and mostly simple spores, and smaller, matt<br />
black pycnidia. M. rhabdogena is similar in appearance and spore characteristics to M. misella.<br />
.
182 BRIAN JOHN COPPINS<br />
but <strong>the</strong> latter has an oHvaceous, K+ violet hymenium (fuscous pigment lacking) and its<br />
mesoconidia are borne in stalked pycnidia. Diminutive, endoxylic forms <strong>of</strong> M. denigrata can be<br />
distinguished by <strong>the</strong> absence <strong>of</strong> a distinct (fuscous) epi<strong>the</strong>cium, hymenium reacting K+ violet,<br />
C+ violet (due to olivaceous pigment) and, usually, C+ orange-red (gyrophoric acid). The<br />
mesoconidial states <strong>of</strong> M. denigrata and M. rhabdogena are very similar, but M. denigrata has<br />
longer (4-5-7-5 /xm) microconidia, and sometimes has curved macroconidia; macroconidia are<br />
apparently not produced by M. rhabdogena or <strong>the</strong> related M. elachista.<br />
Habitat and distribution: M. rhabdogena appears to be a rare or much overlooked, exclusively<br />
lignicolous species, known so far only from north Norway, mid-Sweden and Karelia. All<br />
collections are on ra<strong>the</strong>r hard conifer lignum, and associated species include Calicium trabinellum,<br />
Cetraria pinastri, Mycocalicium subtile, Parmeliopsis ambigua, P. hyperopia, Xylographa<br />
abietina, andX vitiligo.<br />
Exsiccata: Malme Lick. Suec. 20 (M, S).<br />
37. Micarea stipitata Coppins & P. James<br />
(Fig. 29B;Map21)<br />
in Lichenologist 11: 156 (1979). Type: Scotland, Argyll, Loch Creran, Glasdrum National Nature<br />
Reserve, on Betula, 27 v 1976, L. Tibell & Coppins 2357 (E - holotype!; isotypes: BM!, H!, UPS, hb<br />
Poeh!,hbVezda!).<br />
Thallus and apo<strong>the</strong>cia: more or less identical in appearance and internal anatomy to those <strong>of</strong><br />
M. pycnidiophora (q.v.).<br />
Pycnidia always present numerous and conspicuous, borne on distinct whitish stalks (pycnidiophores)<br />
which may be simple, bifurcate or to 5-branched, mainly 400-800 /xm tall and<br />
60-100(-150) fxm diam, <strong>of</strong>ten with small irregular clusters <strong>of</strong> ± superficially encapsulated algae<br />
(small granular-areolae). Stalk tissue composed <strong>of</strong> interwoven hyphae, c. 1-1-5 /xm wide, which<br />
± separate in K. Pycnidia innate in <strong>the</strong> apex or apices <strong>of</strong> pycnidiophores, ± globose or doliiform,<br />
c. 60-90 fxm diam. Conidiogenous cells ± cylindrical, 7-11x1-4-2 /xm. Conidia (mesoconidia)<br />
cylindrical or narrowly ellipsoid, eguttulate, 6-8x1-1-8 fxm.<br />
Chemistry: All parts K- , C- , KC-<br />
, PD-<br />
; no substances detected by t. I.e.<br />
Observations: See under <strong>the</strong> related species M. pycnidiophora for <strong>the</strong> differences between<br />
<strong>the</strong>m and for comparisons with o<strong>the</strong>r similar species.<br />
Habitat and distribution: M. stipitata is characteristic <strong>of</strong> acid, <strong>of</strong>ten leached, bark (or<br />
overgrowing bryophytes <strong>the</strong>reon) on trees in undisturbed woodlands in areas with a high<br />
rainfall. It occurs in communities referable (or closely akin) to <strong>the</strong> Parmelietum laevigatae on <strong>the</strong><br />
trunks <strong>of</strong> Betula, Quercus, and, less frequently, Alnus, Pinus, Abies, and Pseudotsuga. It has<br />
once been found on mossy rocks.<br />
The <strong>British</strong> distribution <strong>of</strong> M. stipitata is a distinctly more nor<strong>the</strong>rn and western, as well as<br />
more oceanic, than that <strong>of</strong> M. pycnidiophora (see Map 21). It has not yet been found in<br />
south-west England but presumably occurs <strong>the</strong>re. Although known from <strong>the</strong> Azores and <strong>the</strong><br />
Canary Islands it has not been found elsewhere in Europe, but should be looked for in <strong>the</strong><br />
hyperoceanic woodlands <strong>of</strong>, for example, south-west Norway and France (Bretagne).<br />
Additional information: for additional illustrations and some fur<strong>the</strong>r discussion see Coppins &<br />
James (1979).<br />
38. Micarea subleprosula (Vezda) Vezda<br />
(Figs 30, 53-54; Map 25)<br />
in Vezda & V. Wirth in Folia geobot. phytotax, Praha 11: 101 (1976). - Bacidia subleprosula Vezda in<br />
Preslia 33: 366 (1961). Type: Czechoslovakia, Bohemia, Sudety, Krkonose, Mumlava valley near<br />
Harrachov, 900 m, over decaying mosses on granitic rocks, 1960, A. Vezda (E - isotype! [t.l.c:<br />
alectorialicacid]).<br />
^
LICHEN GENUS MICAREA IN EUROPE 183<br />
Thallus: As for M. leprosula {q.v.) but with a different chemistry (see below).<br />
Apo<strong>the</strong>cia sometimes (? <strong>of</strong>ten) absent, immarginate, ± globose and constricted at <strong>the</strong> base,<br />
sometimes irregularly aggregated or tuberculate, grey-brown, brownish black, or black and<br />
<strong>of</strong>ten with bluish tinge, 0-2-0-8 mm diam. Hymenium c. 65-90 /xm tall, ± hyaline but upper part<br />
oHvaceous or dilute reddish brown, K— , HNO3-I- reddish. Asci clavate, c. 60-70x19-22 /am.<br />
Spores fusiform, <strong>of</strong>ten slightly curved, 3-7(-9)-septate, (35-)40-50(-60) /xm. Paraphyses<br />
numerous, branched and <strong>of</strong>ten anastomosing, 1-1-5 /itm wide; apices <strong>of</strong>ten more richly<br />
branched, not or only slightly incrassate (to 1-8 /xm). Hypo<strong>the</strong>ciumc. 120-200 /xm tall, ± hyahne<br />
or dull straw, but <strong>of</strong>ten mottled dilute fuscous-brown (K-) in <strong>the</strong> upper part; hyphae<br />
interwoven, but becoming outwardly orientated towards <strong>the</strong> hymenium and excipulum, c.<br />
0-8-1 -5 /xm; ascogenous hyphae with swollen cells, c. 2-6 fxm wide. Excipulum distinct in young<br />
apo<strong>the</strong>cia, but reflexed and obscured in old apo<strong>the</strong>cia, hyaline or dilute fuscous-brown in part;<br />
hyphae radiating, branched and anastomosing, c. 1-1-5 jxm wide.<br />
Pycnidia not found.<br />
Chemistry: Thallus K— , C+<br />
acid plus two unidentified accessory substances.<br />
red, PD-I- yellow; sections <strong>of</strong> apo<strong>the</strong>cia C— ; t.l.c: alectorialic<br />
Observations: Micarea subleprosula is closely allied to M. leprosula and sterile thalli <strong>of</strong> <strong>the</strong> two<br />
species can only be distinguished by <strong>the</strong>ir reactions with PD, or by t.l.c. See under M. leprosula<br />
for fur<strong>the</strong>r discussion.<br />
For additional illustrations <strong>of</strong> A/, subleprosula see Vezda (1961).<br />
Habitat and distribution: M. subleprosula grows over mosses on rocks in much <strong>the</strong> same<br />
habitats in which M. leprosula is mostly commonly found. It was originally described from <strong>the</strong><br />
Sudety mountains <strong>of</strong> Czechoslovakia, where it occurred at an altitude <strong>of</strong> 900 m, and I have seen<br />
only two additional specimens: from Sweden (Varmland) at 200 m; and Wales (Snowdonia) at c.<br />
760 m. It may have been overlooked for M. leprosula. However, I have tested numerous<br />
individual thalli personally ga<strong>the</strong>red in <strong>the</strong> field (mainly in Scotland) during <strong>the</strong> last six years,<br />
and all <strong>of</strong> <strong>the</strong>se were Pd+ red and referable to M. leprosula.<br />
39. Micarea subnigrata (Nyl.) Coppins & Kilias<br />
(Figs. 31A, 50: Map 22)<br />
in Kilias in Herzogia 5: 391 (1980). - Lecidea subnigrata Nyl. in Flora, Jena 49: 370 (1866). - Lecidea<br />
denigrata* [subsp.] subnigrata (Nyl.) Crombie, Lich. Brit.: 70 (1870). - Catillaria subnigrata (Nyl.)<br />
Herre in Proc. Wash. Acad. Sci. 12: 94 (1910). Type: Wales, Merioneth, Cader Idris, 1866, W. A.<br />
Leighton (H-NYL 19136 -lectotype!, sel. Kilias (/oc. dr.); BMexK-isolectotype!).<br />
Lecidea confusula Nyl. in Flora, Jena 55: 360 (1872). - Micarea confusuta (Nyl.) Hedl. in Bih. K. svenska<br />
VetenskAkad. Handl. Ill, 18 (3): 76, 86 (1892). Type: Scotland, East Perthshire, Blair Atholl, Craig<br />
Tulloch, on mica-schist, 1871, Crombie (H-NYL 20191 -lectotype!; BM-isolectotype!).<br />
Thallus effuse, widespreading or as small patches amongst o<strong>the</strong>r lichens, <strong>of</strong> scattered to<br />
confluent or clustered, irregularly convex to subglobose areolae. Areolae 0-08-0-45 mm diam<br />
(usually at <strong>the</strong>ir largest when adjoining apo<strong>the</strong>cia), dark grey-brown, matt or slightly glossy; in<br />
section with a hyaline amorphous covering layer c. 3-10 jxm thick, and outermost hyphae <strong>of</strong>ten<br />
pale brown (K-); a white medulla <strong>of</strong>ten developed in large areolae. Phycobiont mvc^nQOid, cells<br />
c. 4-7 /xm diam.<br />
Apo<strong>the</strong>cia usually numerous, scattered to crowded and confluent, immarginate, at first adnate<br />
and slightly convex, soon becoming convex-hemispherical and occasionally tuberculate, dark<br />
brown (reddish brown when wet) to brownish black, matt or slightly glossy (especially when<br />
young), 0-2-0-6 mm diam, or up to c. 1 mm diam when tuberculate. Hymenium 35-38 /am tall,<br />
hyaline but upper c. 10-12 fxm (epi<strong>the</strong>cium) fuscous brown, K-, HNO3-. Asci clavate<br />
33-35x10-14 /xm. Spores ellipsoid (0-)l -septate, 8-12x4-5 /am. Paraphyses numerous,<br />
branched and anastomosing, (l-)l-3-l-7 /xm wide, sometimes, gradually widening to 2-5 fxm<br />
towards apices; hyaline throughout (epi<strong>the</strong>cial pigment confined to gel-matrix). Hypo<strong>the</strong>cium
184 BRIAN JOHN COPPINS<br />
45-160 ixm tall, hyaline; hyphae interwoven, some becoming vertically orientated towards <strong>the</strong><br />
hymenium, c. 1-2-1 -8 /xm wide, intermixed with ± inflated, short-celled ascogenous hyphae c.<br />
2-5-5 fjcm wide. Excipulum well developed, 25-45 yam wide, dilute fuscous brown; hyphae<br />
radiating, branched and anastomosing, c. 1-3-1-7 jxm wide.<br />
Pycnidia usually present, immersed within areolae with ostioles visible (x50) as minute pores<br />
surrounded by a thin, <strong>of</strong>ten slightly raised, dark brown rim; in section with a simple circular or<br />
flask-shaped cavity, walls hyaline, or pale brown around <strong>the</strong> ostiole; <strong>of</strong> two types: (a) c. 80-200<br />
/xm diam; conidiogenous cells ampulliform to subcylindrical, some with 1-2 percurrent proliferations,<br />
4-10 X 1-5-2-5 /xm, sometimes with swollen base to 4 /xm wide; conida (macroconidia)<br />
helicoid with 2-3 spirals, <strong>of</strong>ten appearing 3-5-septate, overall size 7-10x5-6 /xm, filaments<br />
1-8-2 /xm wide (b) c. 40-100 /xm diam; conidiogenous cells ± cylindrical or ampulliform,<br />
4-lOx 1-1 -4 /xm, sometimes with swollen base to 3 /xm wide; conidia {microconidid) cylindrical,<br />
4-6-6x0-8-1 /xm.<br />
Chemistry: Thallus K— , C-, PD — ; sections <strong>of</strong> apo<strong>the</strong>cia C-; t.l.c: no substances.<br />
Observations: Micarea subnigrata is characterized by its dark grey-brown, verrucose-areolate<br />
thallus, dark brown apo<strong>the</strong>cia, fuscous brown epi<strong>the</strong>cium and excipulum, colourless hypo<strong>the</strong>cium,<br />
and 0-1-septate, eUipsoid spores. However, its most outstanding and characteristic<br />
feature is <strong>the</strong> helicoid shape <strong>of</strong> its macroconidia (Fig. 50A); such conidia are unique in <strong>the</strong> genus<br />
and apparently so in all pycnidial fungi, whe<strong>the</strong>r lichenized or not. In general appearance,<br />
habitat and shape and septation <strong>of</strong> spores M. subnigrata could be confused with M. intrusa, but<br />
Map 22 Micarea intrusa A 1950 onwards A Before 1950 + Micarea subnigrata • 1950 onwards O Before<br />
1950<br />
i
LICHEN GENUS MICAREA IN EUROPE 185<br />
that species has a green epi<strong>the</strong>cium and a large-celled, non-micareoid phycobiont. The little<br />
known M. curvata may be closely related to M. subnigrata, but can be distinguished by <strong>the</strong><br />
fabiform or strongly curved spores and C+ red reaction <strong>of</strong> apo<strong>the</strong>cial sections. With its external<br />
and internal fuscous brown colourations M. subnigrata could be mistaken for a Fuscidea, but<br />
species <strong>of</strong> that genus have broader (c. 1 •7-2-7 /xm) mostly unbranched paraphyses (<strong>the</strong> apical<br />
cell <strong>of</strong> each being clavate or capitate and <strong>of</strong>ten provided with a dark brown 'apical cap'), asci with<br />
a thick, strongly amyloid apical wall, and probably referable to <strong>the</strong> Teloschistes-type (Honegger,<br />
1978), and a protococcoid phycobiont, with thick-walled cells, c. 10-14 ^im diam.<br />
For additional illustrations <strong>of</strong> M. subnigrata see Kilias (1981: 391, 445).<br />
Habitat and distribution: M. subnigrata is confined to hard siliceous rocks in ra<strong>the</strong>r exposed<br />
situations. The communities in which it occurs are attributable to <strong>the</strong> Lecideion tumidae<br />
alliance, especially <strong>the</strong> Lecideetum lithophilae association. The collection {Coppins 8417) from<br />
Glentrool in Kirkcudbrightshire was made from <strong>the</strong> south-facing side <strong>of</strong> a doleritic boulder on a<br />
south-facing slope, at an altitude <strong>of</strong> 135 m; association species were: Cladonia subcervicornis,<br />
Huilia albocaerulescens, H. tuberculosa, Lecanora badia, Lecidea pycnocarpa, Lepraria neglec-<br />
ta, Rhizocarpon obscuratum, R. oederi, Stereocaulon vesuvianum, Trapelia involuta, T. aff.<br />
obtegens, and Andraea sp. From o<strong>the</strong>r <strong>British</strong> collections <strong>the</strong> following can be added: Candela-<br />
riella vitellina, Catillaria chalybeia, Lecanora intricata, L. polytropa, Lecidea fuliginosa, L.<br />
lactea, Lecidella scabra, and Parmelia verruculifera. For information on associated species in its<br />
Norwegian locality see M. intrusa (Table 5).<br />
M. subnigrata is known from scattered localities in upland districts in <strong>the</strong> Scottish highlands,<br />
Galloway, and Wales; recent field studies indicate that it may be far more common in <strong>the</strong>se and<br />
similar areas (e.g. Lake District and Dartmoor) than present records (Map 22) suggest. Outside<br />
Britain it is known only from coastal Norway (Hordaland) and south-west Sweden (Halland).<br />
40. Micarea subviolascens (Magnusson) Coppins, comb. nov.<br />
(Fig. 31B)<br />
Lecidea subviolascens Magnusson in Blyttia 7: 30 (1949). Type: Norway, Hordaland, Granvin, Steinsaethorgen,<br />
on easily wea<strong>the</strong>red schist in an open windy place <strong>of</strong> a small hill, 780 m, ix 1944, J. J. Havaas,<br />
Lich. Exs. Norv. 694 (BG-lectotype!). Topotype material collected in 1949distributed in Havaas, L/c/z.<br />
Exs. Norv. 710 (BG!) and Lich. Norv. Occid. 269 (BG!).<br />
Lecidea assimilata f. aberrans Th. Fr., Lich. Scand. 2: 523 (1874). Type: Norway, Troms, Troms0,<br />
Fl0jfjellet, 1868, Th. M. Fries (UPS-holotype!).<br />
Lecidea assimilata var. hardangeriana Vainio in Acta Soc. Fauna. Fl. fenn. 57 (2): 374 (1934). - Lecidea<br />
assimilata var. hardangeriana Vainio in Havaas in Bergens Mus. Arb. 1909 (1): 29 (1909); nom. nudum<br />
(Art. 32). Type: Norway, Hordaland, Granvin, Sm0reggen, 650 m, 22 viii 1902, J. J. Havaas, Lich. Exs.<br />
Norv. 139 (BG-lectotype! [t.l.c.: no substances]; isolectotypes: BG!, H!).<br />
Thallus effuse, saxicolous, composed <strong>of</strong> ± confluent, white to pale brown, convex, verrucoseareolae<br />
c. 0'l-0-5(-0-8) mm diam; with age <strong>the</strong> crust may thicken (to c. 1 mm) and become<br />
cracked and divided into 'islands' c. 1-2 /xm wide. Areolae in section, sometimes with a hyaline<br />
amorphous covering layer c. 5-7 /xm thick, upper c. 20 /xm <strong>of</strong> areolae <strong>of</strong>ten tinged dilute<br />
olivaceous and Kf+ violet (pigment confined to <strong>the</strong> weak gel-matrix). Thallus hyphae 1-7-2-5<br />
(-3) /xm wide. Phycobiont micareoid, cells 4-7 /xm diam. Cephalodia (?): spaces between<br />
areolae <strong>of</strong>ten filled by black, loose clusters <strong>of</strong> Stigonema.<br />
Apo<strong>the</strong>cia numerous, immarginate, adnate, convex to subglobose, black, matt, 0-3-0-6(-0-8)<br />
mm diam, <strong>of</strong>ten confluent, or forming tuberculate clusters up to 1-2 /xm diam. Hymenium 40-55<br />
/xm tall, dark green (K+ violet, HNO3+ purple-red) above, and below in vertical streaks,<br />
o<strong>the</strong>rwise dilute greenish or ± hyaline. Asci clavate, 38-48x10-15 /xm. Spores ellipsoid,<br />
ovoid-ellipsoid or oblong-ellipsoid, simple, 9-16(-17)x4-5 /xm. Paraphyses numerous, simple<br />
below, but in upper part <strong>of</strong>ten forked or with short lateral branches, l-l-8(-2) /xm wide,<br />
sometimes widening above to 3 /xm; apical walls hyaline although surrounded by densely<br />
pigmented gel-matrix. Hypo<strong>the</strong>cium 150-300 /xm tall, dark purple-brown, K4- purple intensifying,<br />
or upperpart K-l- dark green; all parts HNO3+ purple-red; hyphae interwoven, but ±
186 BRIAN JOHN COPPINS<br />
vertically arranged in upper part, 1 •5-2-5 ^im wide, embedded in densely pigmented gel-matrix;<br />
ascogenous hyphae c. 2-5-5 /xm wide. Excipulum ± distinct in young apo<strong>the</strong>cia, but soon<br />
reflexed, dark purple-brown (K-l- dark green) within, changing to green (K-l- green intensifying)<br />
towards <strong>the</strong> outer edge; hyphae radiating, branched, c. 1-5-2 fxm wide.<br />
Pycnidia: Not found.<br />
Chemistry: Thallus K— , C— , KC— , PD— ; t.l.c: no substances.<br />
Observations: In his protologue, Magnusson allied his new species with <strong>the</strong> Lecidea [Micarea]<br />
sylvicola group. However, Th. M. Fries, 75 years earlier, was probably nearer <strong>the</strong> truth when he<br />
described material <strong>of</strong> M. subviolascens as a form {'aberrans') <strong>of</strong> Lecidea [Micarea] assimilata. In<br />
my opinion M. subviolascens is very closely related to M. assimilata, and <strong>the</strong> reader is referred to<br />
<strong>the</strong> account <strong>of</strong> that species for fur<strong>the</strong>r discussions.<br />
Habitat and distribution: M. subviolascens appears to have a very restricted distribution, being<br />
known only from <strong>the</strong> provinces <strong>of</strong> Troms and Hordaland in Norway. It is <strong>the</strong> only known<br />
saxicolous member <strong>of</strong> <strong>the</strong> M. assimilata group, and occurs on acidic rocks (schists and gneiss) in<br />
ra<strong>the</strong>r open situations. In <strong>the</strong> Hordaland localities it was collected at altitudes <strong>of</strong> 650 m and 795<br />
m. The labels accompanying <strong>the</strong> specimens provide little extra ecological information, and <strong>the</strong><br />
specimens have few associated species, although Pyrenopsis pulvinata and Rhizocarpon obscuratum<br />
have been noted.<br />
Exsiccata: Havaas Lich. Exs. Norv. 139 (BG, H), 694 (BG), 710 (BG). Havaas Lich. Norv. Occid. 269<br />
(BG).<br />
41. Micarea sylvicola (Flotow) Vezda & V. Wirth<br />
(Figs31C,51A;Map23)<br />
in Folia geobot. phytotax, Praha 11: 99 (1976). - Lecidea sylvicola Flotow, Lich. Schles. 171 (1829).<br />
Type: Poland, Schlesien, Flotow, Lich. Exs. 171A (UPS - lectotype!, sel. Hertel (1975: 74); WRSL-<br />
isolectotype!).<br />
Lecidea aggerata Mudd, Man. Br. Lich.: 208 (1861). Type: England, Yorkshire, Battersby, Mudd, Lich.<br />
Brit. 175 (BM - lectotype! ; isolectotypes: E!, H-NYL 14016!, M!, MANCH!).<br />
Lecidea incincta Nyl., Lich. Scand. : 231 (1861). Type: Finland, Satakunta, Kallfjard [Ahlainen: Kellahti],<br />
1859, /i. /. Malmgren (H - holotype!).<br />
Biatora smaragdina ArnoXd'm Verh. zool. bot. Ges. Wien 19: 613 (1869). Type: Italy, Trentino-Alto Adige,<br />
'Melaphyr in Walde unterhalb Bad Razzes am Schiern [Monte Pez]', vii 1867, Arnold (M - holotype!).<br />
Lecidea hellbomii Lahm in Flora, Jena 53: 177 (1870). Type: Sweden, Narke, Vredstorp, Urby, on granitic<br />
rock, 1869, P. J. Hellbom (M- lectotype!).<br />
Lecidea sylvicola f. sublivida Vainio in Medd. Soc. Fauna Fl. fenn. 10: 104 (1883). - Lecidea sylvicola var.<br />
sublivida (Vainio) Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 298 (1934). Type: Finland, Ostrobottnia<br />
kajanensis, Kuhmo,Kylmala, 1877, E. A. Vainio (TUR-VAINIO 25226 -holotype!).<br />
Lecidea hypocyanea Vainio in Acta Soc. Fauna Fl.fenn. 57 (2): 300 (1934), non Stirton (1879). - Lecidea<br />
vainioi Magnusson in Blyttia 7: 31 (1949); nom. nov. Type: Finland, Regio aboensis, Turku, Hirvensalo,<br />
10 vii 1924, E. A. Vainio (TUR-VAINIO 33172 -lectotype!, sel. Hertel (1975: 74)).<br />
Lecidea sylvicola \ar.flotowii Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 297 (1934); nom. inval. (Art. 26).<br />
Thallus effuse, thin and ± smooth or weakly rimose to ra<strong>the</strong>r thick, uneven and coarsely<br />
rimose, or sometimes with irregularly granular or verrucose areolae up to 0-3 mm diam, pale<br />
buff, pale to dark grey or bluish grey; in section without a cortex or hyahne covering layer, but<br />
walls <strong>of</strong> outermost hyphae sometimes greenish, K— , HNO^-f- purple-red. Phycobiont not<br />
micareoid; cells thin-walled, ± globose and c. 5-12 /xm diam, or ellipsoid and up to 15x 10 fxm.<br />
Apo<strong>the</strong>cia numerous, convex-hemispherical from <strong>the</strong> start, <strong>of</strong>ten becoming ± globose or<br />
tuberculate, black and <strong>of</strong>ten with bluish tinge, sometimes dark blue-grey (deep shade forms),<br />
0-2-0-5 diam, or to 1-2 mm when tuberculate. Hymenium 40-60(-70) /xm tall, dilute bright or<br />
sordid aeruginose, but <strong>of</strong>ten darkish aeruginose in upper and lower parts, and in vertical streaks<br />
(due to presence <strong>of</strong> stout pigmented paraphyses), K— or -I- aeruginose intensifying, HNO3-I-<br />
purple-red. Asci cylindrical-clavate, c. 35-45x8-12 /xm. Spores ellipsoid or ovoid, simple (an<br />
occasional 1-septate spore seen in a few collections), (6-)7-10x(2-5-)3-4-5 /am. Paraphyses
LICHEN GENUS MICAREA IN EUROPE 187<br />
ra<strong>the</strong>r scanty, <strong>of</strong> two types: p.p. evenly distributed, irregularly flexuose, simple or sparingly<br />
branched, <strong>of</strong>ten anastomosing, hyaline, thin, 0-7-1 /xm wide, sometimes widening above to 1-8<br />
fim; p.p. fewer in number, as scattered individuals or in small fascicles, straight, simple or<br />
occasionally forked above , stout, coated ± throughout (but most intensely so around <strong>the</strong> apices)<br />
by dark green pigment and appearing c. 2-3 /xm wide, apices sometimes incrassate and up to 4<br />
/Lim wide. Hypo<strong>the</strong>cium 90-200 /xm tall, dark aeruginose or olive-black, K- or + green<br />
intensifying; lower ('core') part, or rarely <strong>the</strong> entire hypo<strong>the</strong>cium, dark purplish brown, K-lpurple<br />
intensifying; all pigments HNO3+ purple-red; hyphae coated with dense dark green<br />
pigment and 2-3 /xm wide (overall), interwoven but becoming vertically orientated towards <strong>the</strong><br />
hymenium and some continuing into it as stout, pigmented paraphyses; ascogenous hyphae<br />
similarly pigmented, with swollen cells c. 3-5 /xm wide. Excipulum not evident, even in sections<br />
<strong>of</strong> young apo<strong>the</strong>cia.<br />
Pycnidia usually present and numerous, immersed, black, 40-200 /xm diam; in section with a<br />
single circular or ovate locule but <strong>of</strong>ten becoming internally convoluted with up to 6 locules, wall<br />
dark green K- , HNOs-f- purple-red in upper part, turning paler below and dilute brownish or ±<br />
hyaline at <strong>the</strong> base; conidiogenous cells irregularly subcylindrical, 5-10 x 1-2-1 -5 /xm, <strong>of</strong>ten with<br />
one or two percurrent proliferations; conidia (mesoconidia) ± cylindrical or sometimes oblongobovoid,<br />
sometimes biguttulate and <strong>of</strong>ten ± constricted in <strong>the</strong> middle, 3-8-6(-6-6)xl-l-7(-2)<br />
/xm.<br />
Chemistry: Thallus K-, C-, KC-, PD-; sections <strong>of</strong> apo<strong>the</strong>cia C-; t.l.c: no substances.<br />
Observations: Micarea sylvicola is characterized by its convex-globose, <strong>of</strong>ten tuberculate,<br />
black apo<strong>the</strong>cia, green or blue-green hymenium, very dark green-black or brown-black<br />
hypo<strong>the</strong>cium (appearing green, or rarely purplish, in K), and ellipsoid to ovoid, simple spores.<br />
The variation in hypo<strong>the</strong>cial pigmentation is similar to that found in such species as M.<br />
assimilata, M. crassipes, and M. melaena. M. tuberculata is ra<strong>the</strong>r similar to M. sylvicola but has<br />
generally smaller apo<strong>the</strong>cia, a more shallow hymenium, narrower, oblong-ovoid spores that are<br />
<strong>of</strong>ten 1-septate, and shorter conidia. Forms <strong>of</strong> M. bauschiana with greenish pigment in <strong>the</strong><br />
hypo<strong>the</strong>cium have been confused with M. sylvicola, however <strong>the</strong> pigment is always dilute and<br />
confined to <strong>the</strong> gel-matrix such that <strong>the</strong> hyphae appear hyaline in K (at x 1000) . The thallus <strong>of</strong> M.<br />
sylvicola is <strong>of</strong>ten provided with a blue-grey tinge due to green-pigmented hyphae near <strong>the</strong><br />
surface; such coloration has not been noticed with <strong>the</strong> thalli <strong>of</strong> M. bauschiana and M.<br />
tuberculata. Ano<strong>the</strong>r species <strong>of</strong>ten confused with M. sylvicola is Lecidea erratica Korber, which<br />
can be distinguished by its thinly marginate young apo<strong>the</strong>cia, and distinct excipulum when<br />
observed in section.<br />
Habitat and distribution: M. sylvicola is mainly found on dry shaded rocks in <strong>the</strong> communities<br />
belonging to Micareetum sylvicolae, but it sometimes occurs in periodically wetter situations on<br />
rock faces, upper sides <strong>of</strong> boulders, and loose stones in woodlands where it may be associated<br />
with such species as Baeomyces rufus, Cystocoleus ebeneus, Fuscidea recensa, Huilia tuberculosa,<br />
Parmelia saxatilis, Rhizocarpon hochstetteri, R. obscuratum, R. oederi, Scoliciosporum<br />
umbrinum, Trapelia involuta, and in one instance (Coed Hafod, Denbigh) Bacidia vezdae. In<br />
addition, it is sometimes found on old fence posts in upland districts.<br />
In <strong>the</strong> <strong>British</strong> Isles M. sylvicola is more confined to upland districts and is less common than M.<br />
bauschiana. It seems to be particularly prevalent in Wales and south-east Scotland, and<br />
curiously rare in western Scotland and Ireland. Fur<strong>the</strong>r field-work is required to establish<br />
whe<strong>the</strong>r or nor its apparent scarcity in <strong>the</strong>se latter areas is real or merely an artefact resulting<br />
from uneven recording. However, my field observations in western Scotland suggest that its<br />
niche in <strong>the</strong> Micareetum sylvicolae is <strong>the</strong>re occupied by M. lutulata. M. sylvicola is widely<br />
distributed in Scandinavia, but rarely found from north <strong>of</strong> about latitude 67°N. Elsewhere in<br />
Europe I have seen specimens from Germany (Hessen and Baden-Wiirttemberg), eastern<br />
Austria, nor<strong>the</strong>rn Italy, south-west Poland and <strong>the</strong> mountain regions <strong>of</strong> Czechoslovakia. From<br />
outside Europe I can confirm its presence in north-eastern North America (New York and<br />
Newfoundland).
188 BRIAN JOHN COPPINS<br />
Map 23 Micarea sylvicola # 1950 onwards O Before 1950<br />
Exsiccata: Arnold Lich. Exs. 409A (BM ex K, M, MANCH, WRSL), 409Bp./7. (BM ex K, M). Flotow<br />
Lich. exs. 171A (WRSL). Johnson Lich. Herb. 434 p.p. (BM). Kav. & Hilitzer Crypt. Cech. 269 (UPS).<br />
Korber Lich. Sel. Germ. 75 (M). Larb. Lich. Caesar. Sarg. 84 (ABD, BM, MANCH). Larb. Lich. Herb.<br />
304, 305 p.p. (BM). Malme Lich. Suec. 199 (M, S). Mudd Lich. Brit. 175 (BM, E, M, MANCH). Norrl. &<br />
Nyl. Herb. Lich. Fenn. 763 (BM, H, TUR, WIS), 764 (BM, H, WIS). Rasanen Lich. Fenn. 612 p.p. (M).<br />
Suza Lich. Bohem. 131 (M). Vezda Lich. Sel. 957, 1341 (BM). Zwackh Lich. Exs. 535, 596 (M), 597<br />
(UPS), 780 (M), 919 (BM, M).<br />
42. Micarea syno<strong>the</strong>oides (Nyl.) Coppins<br />
(Figs . 3 1 D-E , 43D-E ; Map 24)<br />
in Topham & Walker in Lichenologist 14: 67 (1982) - . Lecidea syno<strong>the</strong>oides Nyl . , Lich. Jap. : 63 ( 1 890) . -<br />
Catillaria syno<strong>the</strong>oides (Nyl.) Zahlbr., Cat. lich. univ. 4: 78 (1926). Type: Japan, 'Itjgome' (or<br />
'Itchigome'), on lignum, 1879, E. Almquist (H-NYL 19101 - lectotype!; S - isolectotype!).<br />
Thallus effuse, <strong>of</strong> scattered to coherent, or clustered, granular areolae. Areolae c. 20-70 /am<br />
diam, dull grey-green (never whitish) to dark olivaceous or blackish, appearing ± gelatinous<br />
when wet, usually ± discrete but <strong>of</strong>ten coalescing to form a thin uneven crust; a thin white<br />
prothallus sometimes visible between scattered areolae. Areolae in section without an amorphous<br />
covering layer; outer hyphae <strong>of</strong>ten with olivaceous or brownish, K-l- violet walls.<br />
Phycobiont micareoid, cells c. A-1 fxva.<br />
Apo<strong>the</strong>cia immarginate, convex-hemispherical to ± globose, <strong>of</strong>ten tuberculate, grey-black or<br />
brown-black, matt, sometimes grey (shade forms), 0- 1-0-3 mm diam, or up to 0-5 mm diam
LICHEN GENUS MICAREA IN EUROPE 189<br />
when tuberculate. Hymenium 35-45 /xm tall, dilute olivaceous or dilute olive-brown, K+ violet:<br />
pigment confined to gel-matrix. Asci clavate, 30-40x9-5-12 jxm. Spores ± acicular or rodshaped,<br />
curved or ± straight, l-7(-ll)-septate, 14^35(-43)xl-8-2-5(-3) y,m. Paraphyses<br />
numerous, branched, sometimes anastomosing, 0-8-1 /xm wide, sometimes widening to 1-5 /am<br />
towards <strong>the</strong>ir apices; apical walls hyaline. Hypo<strong>the</strong>cium 60-100 jxm tall, hyaline, or dilute<br />
olivaceous and <strong>the</strong>n Kf+ violet; hyphae hyaline, 1-2 ^im wide, interwoven or some ± vertically<br />
orientated in <strong>the</strong> upper part; intermixed with short-celled ascogenous hyphae, c. 2-4 ^im wide.<br />
Excipulum indistinct, evident in sections as a narrow, reflexed, non-amyloid lateral border to<br />
<strong>the</strong> hymenium, hyaline or pale olivaceous (<strong>the</strong>n K+ violet), varying from paler to darker than<br />
<strong>the</strong> hymenium; hyphae hyahne, radiating branched and anastomosing, c. 1 /xm wide.<br />
Pycnidia frequent but inconspicuous, immersed within areolae, or emergent to sessile,<br />
whitish to grey-black; walls dull olivaceous (pigment <strong>of</strong>ten more intense around <strong>the</strong> ostioles),<br />
K+ violet. Pycnidia <strong>of</strong> two types: (a) 60-120 /am diam, immersed, or emergent with gaping<br />
ostioles; conidia {mesoconidia) ± cylindrical to fusiform, 4-5-6xl-2-l-5 /xm; (b) 30-40 /Ltm<br />
diam, immersed to sessile, but ostioles not gaping; conidia (microconidia) short-cylindrical,<br />
3-8-4-8x0-8-l/xm.<br />
Chemistry: Thallus C— , K— , PD— ; sections <strong>of</strong> thallus and apo<strong>the</strong>cia C- (but olivaceous parts<br />
C-l- violet due to pigment); t.l.c: no substances detected.<br />
Observations: Micarea syno<strong>the</strong>oides is characterized by its ± acicular or rod-shaped spores,<br />
K-l- violet hymenium, and olivaceous thallus which is j. gelaimous when wet. M. globulosella is<br />
very similar, but differs in having a thallus <strong>of</strong> whitish or grey, somewhat larger areolae<br />
containing gyrophoric acid. M. nitschkeana differs in <strong>the</strong> same ways, and in addition has shorter<br />
(rarely over 17 jxm) spores and longer microconidia. M. globulosella and M. syno<strong>the</strong>oides have<br />
been confused with Bacidia beckhausii (see p. 196) which has similar spores, pigmentation and<br />
paraphyses, but can be distinguished by its large-celled (c. 8-14 fxm diam) phycobiont,<br />
excipulum <strong>of</strong> coherent (in K) hyphae, smaller conidia (one type only, c. 2-8-3-5X 1-1-4 /xm),<br />
<strong>of</strong>ten pruinose apo<strong>the</strong>cia, and usual occurrence on less acidic bark (especially <strong>of</strong> Acer, Fraxinus,<br />
Populus, and Ulmus), although it does occasionally occur on Quercus trunks and more rarely on<br />
lignum.<br />
Of <strong>the</strong> four Japanese specimens, two are on bark and two (type material) are on lignum. They<br />
all have ra<strong>the</strong>r siiprt spores (14-26 /xm), but appear to agree in most o<strong>the</strong>r respects with<br />
collections from Britain and <strong>the</strong> Atlantic islands. Unfortunately I was unable to find mesoconi-<br />
dia in <strong>the</strong> Japanese specimens. For <strong>the</strong> time being, I consider <strong>the</strong> Japanese and Atlantic<br />
populations as conspecific, especially as <strong>the</strong> <strong>British</strong> collections exhibit considerable variation in<br />
spore septation and length. Fur<strong>the</strong>r collections from Japan are required to finally establish <strong>the</strong><br />
relationship between <strong>the</strong> two populations.<br />
Habitat and distribution: M. syno<strong>the</strong>oides is an oceanic species mostly found in wooded valleys<br />
on bark or over bryophytes, on <strong>the</strong> trunks <strong>of</strong>, for example, Alnus, Betula, Quercus, Larix,<br />
Pinus, and Pseudotsuga, in communities in, or related to, <strong>the</strong> Parmelion laevigatae alliance.<br />
<strong>British</strong> collections include <strong>the</strong> following associated lichens: Bryoria fuscescens, Chrysothrix<br />
candelaris, Cladonia coniocraea, C. squamosa, Haematomma elatinum, Hypogymnia physodes,<br />
Lecanactis abietina, Lecidea icmalea, Micarea alabastrites, M. cinerea, M. peliocarpa, M.<br />
stipitata, Mycoblastus sterilis, Ochrolechia androgyna, Parmelia saxatilis, Pertusaria amara,<br />
Platismatia glauca, Sphaerophorus globosus, Trapelia corticola in ed., and Usnea subftoridana.<br />
The single <strong>British</strong> collection on lignum {Coppins 2299) was on a rotting trunk <strong>of</strong> Pinus. In <strong>the</strong><br />
Canary Islands it occurs on Erica arborea in a photophobus and hydrophilic community<br />
composed <strong>of</strong> crustose lichens with Atlantic distributions (Topham & Walker, loc. cit.). In <strong>the</strong><br />
Azores it occurs in a more or less identical community on Cryptomeria.<br />
Apart from <strong>the</strong> type locality in Japan, it is known only from western Britain, <strong>the</strong> Azores, and<br />
<strong>the</strong> Canary Islands (Tenerife). It should be sought for in suitable habitats in o<strong>the</strong>r oceanic parts<br />
<strong>of</strong> Europe (e.g. south-west Norway, Bretagne, <strong>the</strong> Pyrenees, and Portugal).
190 BRIAN JOHN COPPINS<br />
Map 24 Micarea globulosella ® 1950 onwards + Micarea syno<strong>the</strong>oides # 1950 onwards O Before 1950<br />
43. Micarea ternaria (Nyl.) Vezda<br />
(Figs 32B, 52; Map 25)<br />
Sched. Lich. Sel. Exsicc. 858 (1970). - Lecidea sabuletorum f. ternaria Nyl. in Not. Sallsk Fauna Fl. fenn.<br />
Forhandl. 8: 151 (1866). - Lecidea ternaria (Nyl.) Nyl. in Flora, Jena 60: 232 (1877). Type: USSR,<br />
Lapponia murmanica, Kola Peninsula, 'Olenji', 1861, N. I. Fellman (H-NYL 18682 -holotype! [t.l.c:<br />
no substances]).<br />
Thallus effuse, muscicolous, composed <strong>of</strong> scattered to confluent, convex to irregularly<br />
subglobose areolae [? or saxicolous and obsolete -see 'observations']. -Areolae cream-white to<br />
ash-grey, c. 80-300 /xm diam; in section with hyahne amorphous covering layer c. 7-10 /xm tall,<br />
and outermost hyphae with greenish (K— , HNO3-I- red) walls. Phycobiont micareoid, cells c.<br />
4—7)Ltm diam.<br />
Apo<strong>the</strong>cia numerous, black, matt or slightly glossy, 0-15-1 mm diam; at first (below 0-3 mm<br />
diam) ± globose or turbinate, later expanding to become broadly convex and ± adnate,<br />
sometimes faintly marginate. Hymenium c. 60-70 /xm tall, merging imperceptibly into <strong>the</strong><br />
hypo<strong>the</strong>cium; upper c. 10-15 ixm (epi<strong>the</strong>cium) dark aeruginose-green, K-, HNO3+ purplered;<br />
remaining (lower) part ± hyaline with dilute green vertical streaks. Asci clavate. Spores<br />
fusiform, sometimes slightly curved, (0-)l-3-septate, 13-22(-24)x 3-5-5 /am. Paraphyses<br />
numerous, sparingly branched below but richly branched in epi<strong>the</strong>cium, <strong>of</strong>ten anastomosing, c.<br />
1-1-8 /Am wide; apices <strong>of</strong>ten slightly incrassate and surrounded by dense pigment, and up to 3<br />
fxm wide. Hypo<strong>the</strong>cium up to 380 /am tall; upper third to half, dilute olivaceous or dilute dull
LICHEN GENUS MICAREA IN EUROPE 191<br />
brown; lower part ± hyaline; hyphae hyaline c. 0-7-1 -5 fim wide, interwoven or becoming<br />
vertically or outwardly orientated towards <strong>the</strong> hymenium and excipulum; hyphae in upper part<br />
<strong>of</strong> hypo<strong>the</strong>cium intermixed with inflated, short-celled ascogenous hyphae. Excipulum usually<br />
distinct, hyaline, or tinged aeruginose or olive-green in parts; hyphae radiating, branched and<br />
anastomosing, 0-7-1 -7 /xm wide.<br />
Pycnidia ra<strong>the</strong>r numerous, sessile, black, 100-140(-200) /xm diam, ostioles <strong>of</strong>ten gaping;<br />
walls dark olive-green (K— , HNO3-I- red) above and at <strong>the</strong> sides, but becoming hyaline towards<br />
<strong>the</strong> base. Conidia (mesoconidia) cylindrical or oblong-ellipsoid, sometimes faintly biguttulate<br />
and slightly constricted in <strong>the</strong> middle, 4-6-6-3x1-2-1-7 /itm. Macro- or microconidial states not<br />
found.<br />
Chemistry: Thallus K-, C— , PD— ; sections <strong>of</strong> apo<strong>the</strong>cia C-; t.l.c: no substances.<br />
Note: The above description is based largely on Alaskan specimens (Thomson 9188).<br />
Observations: M. ternaria is very close to M. lignaria but differs in having shorter spores that<br />
are never more than 3-septate, and in completely lacking lichen substances. In addition its<br />
apo<strong>the</strong>cia have a more flattened appearance (sometimes with a faint marginal rim), and a ±<br />
clearly defined excipulum when viewed in vertical section. It is superficially similar to dark forms<br />
<strong>of</strong> M. peliocarpa but that species contains gyrophoric acid (thallus and apo<strong>the</strong>cia in section C-lorange-red),<br />
has a hyaline hypo<strong>the</strong>cium, and is not known to have pycnidia which contain<br />
mesoconidia.<br />
Having found most herbarium specimens named 'ternaria' to belong to M. lignaria or M.<br />
peliocarpa I suspected <strong>the</strong> name to be a synonym <strong>of</strong> one <strong>of</strong> <strong>the</strong>se two species. The holotype <strong>of</strong><br />
Map 25 Micarea subleprosula (§) 1950 onwards -I- Micarea cf. ternaria # 1950 onwards O Before 1950
192 BRIAN JOHN COPPINS<br />
Lecidea milliaria f. ternaria is very fragmentary and my first impression was that it was a<br />
diminutive specimen <strong>of</strong> M. lignaria, even though I failed to obtain a positive reaction with PD.<br />
This opinion was changed by my subsequent examination <strong>of</strong> <strong>the</strong> Alaskan material which is<br />
identical to <strong>the</strong> holotype in all respects but larger and in better condition.<br />
Several problematical collections from <strong>the</strong> <strong>British</strong> Isles (Shetland Islands and western<br />
Ireland) are provisionally referred to M. ternaria. They were all collected in coastal districts on<br />
hard siliceous rocks and have a ± obsolete thallus (PD— in microscopical preparations), ra<strong>the</strong>r<br />
flattened apo<strong>the</strong>cia, and 0-3-septate spores. The specimen from Fair Isle has pycnidia,<br />
containing mesoconidia5-2-6-8xl-2-l-8/x,m (Fig. 52C). The discovery <strong>of</strong> an arctic species, such<br />
as M. ternaria, in coastal Britain is not without precedent. Examples are Lecanora straminea<br />
found in <strong>the</strong> Shetland Islands, Flannan Islands, and St Kilda, and Bacidia subfuscula found on<br />
North Rona, <strong>the</strong> Fame Islands, and as far south as <strong>the</strong> north coast <strong>of</strong> Norfolk and <strong>the</strong> Scilly Isles.<br />
A better understanding <strong>of</strong> <strong>the</strong> variability in habit and habitat <strong>of</strong> M. ternaria in <strong>the</strong> arctic should<br />
help to clarify <strong>the</strong> taxonomic relationships between <strong>the</strong> arctic and <strong>British</strong> populations. In<br />
addition, relevant material should be sought in coastal Norway.<br />
Habitat and distribution: On mosses and plant debris on <strong>the</strong> ground in coastal regions <strong>of</strong> arctic<br />
Europe and North America (nor<strong>the</strong>rn Alaska). The Alaskan material {Thomson 9188) was<br />
growing with Micarea turfosa and Siphula ceratites. Sou<strong>the</strong>rn populations on rocks may be<br />
represented in <strong>the</strong> <strong>British</strong> Isles (see 'observations' above).<br />
44. Micarea tuberculata (Sommerf.) R. Anderson<br />
(Figs32A,34,51B;Map26)<br />
in Bryologist 11: 46 (1974). - Lecidea tuberculata Sommerf., Suppl. Fl. Lapp.: 160 (1826). Type:<br />
Norway, Nordland, Saltdalen, Fiskevaagmollen, iii 1822, S. C. Sommerfelt (O - lectotype!; UPS -<br />
isolectotype!).<br />
Lecidea latens Taylor in Mackay, Fl. Hib. 2: 259 (1836). Type: Ireland, Wicklow, The Dargle, T. Taylor<br />
(BM- lectotype!).<br />
Lecidea botryocarpa Nyl. in Flora, Jena 48: 603 (1865). Type: USSR, Karelskaya ASSR, Karelia<br />
onegensis, 'ad Onegam, Kapselka, 1863, T. Simming (H-NYL 10766 -holotype!).<br />
Lecidea subinfidula Nyl. in Flora, Jena 52: 295 (1869). Type: Finland, Lapponia enontekiensis, Naimakka,<br />
29 viii 1867, J. P. Norrlin 656 (H - lectotype!; UPS - isolectotype!).<br />
Lecidea tuberculata var. scandinavica Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 309 (1934). Type: Finland,<br />
Tavastia australis, Lammi, Evo, Lapinkallio, 1866, Norrlin (H-NYL 10767 - lectotype!).<br />
Thallus effuse, minutely scurfy-granular, or sometimes forming an irregularly rimose crust up<br />
to 0-2 mm thick, more rarely forming irregularly verrucose areolae c. 0-06-0-2(-0-3) mm diam;<br />
greenish buff, pale buff or greenish white; thin wefts <strong>of</strong> white prothalline hyphae sometimes<br />
visible. Phycobiont not micareoid; cells ± globose, thin-walled, 5-10(-12) /xm diam, or<br />
irregularly ellipsoid and up to 15x7 /xm.<br />
Apo<strong>the</strong>cia numerous, convex-hemispherical and immarginate from <strong>the</strong> start, <strong>of</strong>ten becoming<br />
± globose or tuberculate, black and <strong>of</strong>ten with bluish tinge, matt, 0-16-0-3(-0-4) mm diam, or<br />
0-24-0-55 mm diam when tuberculate. Hymenium 30-35 jxm tall, dilute green or aeruginose,<br />
K- or + intensifying, HNO3+ purple-red, <strong>of</strong>ten with darker vertical streaks due to <strong>the</strong> presence<br />
<strong>of</strong> stout, pigmented paraphyses. Asci cylindrical-clavate, 25-30x7-9 /xm. Spores oblong-ovoid<br />
or oblong-ellipsoid, 0-1-septate, 5-5-8(-9)xl-5-2-5 /xm. Fflrap/iy:?^^ ra<strong>the</strong>r scanty, <strong>of</strong> two types<br />
(Fig. 34): p.p. evenly distributed, irregularly flexuose, simple or sparingly branched, <strong>of</strong>ten<br />
anastomosing, thin, 1-T5 /xm wide, sometimes widening to 1-7 /xm above, walls hyaline<br />
throughout and without adhering pigment; /?./?. fewer in number, as scattered individuals or in<br />
small fascicles, straight, simple or occasionally forked above, stout, coated ± throughout by<br />
dark greenish pigment and appearing c. 2-3 /u-m wide, apices sometimes ± incrassate and up to 4<br />
/xm wide (including pigment). Hypo<strong>the</strong>cium c. 80-115 /xm tall, aeruginose- or olive-black, K-laeruginose<br />
intensifying; hyphae coated with dense dark green pigment and 2-3 /xm wide<br />
(overall), interwoven but becoming vertically orientated towards <strong>the</strong> hymenium and some
LICHEN GENUS MICAREA IN EUROPE 193<br />
continuing into it as stout, pigmented paraphyses; ascogenous hyphae similarly pigmented, with<br />
swollen cells c. 3-5 fxm wide. Excipulum not evident, even in sections <strong>of</strong> young apo<strong>the</strong>cia.<br />
Pycnidia <strong>of</strong>ten present, ± immersed, black, 60-120 /u,m diam, ostioles <strong>of</strong>ten gaping; walls dark<br />
aeruginose, K— , HNO3+ red; conidiogenous cells slender, cylindrical, 5-10x1-1-5 /u,m, <strong>of</strong>ten<br />
with a swollen, sometimes pigmented, base up to 3-2 jxm wide, and sometimes with one or two<br />
percurrent proliferations; conidia (mesoconidia) ± cylindrical, sometimes faintly biguttulate,<br />
3-4-3 X 1-1 -4 /xm.<br />
Chemistry: Thallus K— , C— , KC— , PD-; sections <strong>of</strong> apo<strong>the</strong>cia C—; t.l.c: no substances.<br />
Observations: Micarea tuberculata is characterized by its ra<strong>the</strong>r small, markedly convex, <strong>of</strong>ten<br />
tuberculate, blackish apo<strong>the</strong>cia, blue-green hymenium, aeruginose-black hypo<strong>the</strong>cium and<br />
small, ovoid-oblong, 0-1-septate spores. The relative proportion <strong>of</strong> simple to 1-septate spores<br />
can vary greatly between collections, and in some specimens no septate spores can be found. M.<br />
tuberculata has <strong>of</strong>ten been confused with forms <strong>of</strong> M. sylvicola with small, immature spores, but<br />
such spores are always ellipsoid or ovoid-ellipsoid, and quite distinct from <strong>the</strong> predominantly<br />
oblong-ovoid spores <strong>of</strong> M. tuberculata. If pycnidia are present <strong>the</strong> two species can be separated<br />
on <strong>the</strong> size <strong>of</strong> <strong>the</strong>ir conidia, which are significantly longer (c. 4-6)u,m) in M. sylvicola. M. olivacea<br />
(q.v.) is superficially similar to M. tuberculata, but has slightly larger spores, shorter conidiogenous<br />
cells, and a micareoid phycobiont. Also <strong>of</strong> similar appearance is Psilolechia clavulifera<br />
(p. 201), but it has smaller spores, a pale hypo<strong>the</strong>cium, numerous, stout, non-pigmented<br />
paraphyses, and a very different phycobiont.<br />
Map 26 Micarea tuberculata • 1950 onwards O Before 1950
194 BRIAN JOHN COPPINS<br />
Habitat and distribution: M. tuberculata is found on rocks, stones, and tree roots, etc. in dry,<br />
sheltered underhangs, and is a faithful member <strong>of</strong> <strong>the</strong> Micareetum sylvicolae. In <strong>the</strong> <strong>British</strong> Isles<br />
it has scattered localities in <strong>the</strong> north and west; although not yet known from <strong>the</strong> mainland in<br />
England and Wales it is expected to occur in those areas in suitable terrain (e.g. <strong>the</strong> Lake<br />
District, Dartmoor and Snowdonia). It is undoubtedly much overlooked, but appears to be<br />
genuinely rarer than M. bauschiana and M. sylvicola, with which it <strong>of</strong>ten occurs.<br />
It appears to be widely distributed in Fennoscandia where it is known from as far north as<br />
Nordland in Norway and Lycksele Lappmark in Sweden. It seems to be rare elsewhere in<br />
Europe, but I have seen material from sou<strong>the</strong>rn Germany, nor<strong>the</strong>rn Italy, and <strong>the</strong> Tatra<br />
mountains <strong>of</strong> Czechoslovakia.<br />
Exsiccata: Arnold Lich. Exs. 1057 (BM ex K, M). Larb. Lich. Herb. 221 p.p. (BM). Rabenh. Lich. Eur.<br />
648 p.p. (BM). Rasanen Lich. Fenn. 512 (BM, BM ex K), 672 p.p. (BM, LD-mixed with M. lutulata,<br />
M-mixed with M. sylvicola).<br />
\<br />
45. Micarea turfosa (Massal.) Du Rietz<br />
(Fig. 33; Map 27)<br />
in Svensk hot. Tidskr. 17: 94 (1923). - Biatora turfosa Massal., Ric. Lich. Crost.: 128 (1852). Type:<br />
specimen unlocalized but probably from Sudety (Sudeten Mountains) in SW Poland, 'Flotow Lich. exs.<br />
130! hb. Flotow' (VER-holotype!).<br />
Biatora turfosa* verrucula Norman in K. nor. Vidensk. Selsk. Skr. 5: 353 (1868). - Lecidea verrucula<br />
(Norman) Th. Fr., Lich. Scand. 2: 523 (1874). - Micarea verrucula (Norman) Hedl. in Bih. K. svenska<br />
VetenskAkad. Handl. Ill, 18 (3): 84, 95 (1892). Type: Norway, Nordland, 'supra Heminghyt convallis<br />
Bejeren',/. M. Norman (-lectotype!; isolectotypes: LD!, M!, O!).<br />
Thallus effuse but <strong>of</strong>ten forming rounded patches about 3-5 cm diam, thin, ± uneven but not<br />
forming distinct areolae or goniocyst-like granules, blackish grey or brown-black but shaded<br />
portions sometimes dull grey-green, <strong>of</strong>ten appearing ± gelatinous when moist. Thallus in<br />
section up to 70 /xm thick, ecorticate and without an amorphous covering layer; outermost<br />
hyphae 2-4 /am wide with walls thickened by dark green pigment, K-, HNO3+ red; internal<br />
hyphae hyaline, c. 1-1-5 /xm wide. Phycobiont micareoid, cells 4-7 jxm diam.<br />
Apo<strong>the</strong>cia numerous and <strong>of</strong>ten confluent, immarginate, convex to ± globose, black, matt or<br />
slightly glossy, rarely brown (shade forms), 0-15-0-3(-0-4) mm diam. Hymenium 35-50 ^im tall;<br />
upper part (epi<strong>the</strong>cium) sordid green, K— , HNOs-l- red, or sometimes brownish in part; middle<br />
part dilute aeruginose or olivaceous; lower part dilute reddish brown and merging ± impercep-<br />
tibly into <strong>the</strong> hypo<strong>the</strong>cium. Asci clavate, 35-50 x 11-12 /xm. Spores oblong-ellipsoid to fusiform,<br />
sometimes slightly curved, simple or 1-3-septate, (10-)12-21(-25)x(3-5-)4-5 /itm. Paraphyses<br />
numerous, branched and sometimes anastomosing, c. 1-1-5 /xm wide in mid-hymenium but<br />
upper 5-15(-25) fxm thickened with dark greenish pigment and <strong>the</strong>n l-5-2-5(-3) jxm wide.<br />
Hypo<strong>the</strong>cium c. 70-140 /xm tall, mottled reddish brown, K-, HNO3- or turning orange-brown<br />
(never with purple tinge); hyphae interwoven but becoming vertically orientated towards <strong>the</strong><br />
hymenium, hyaline or sometimes loosely coated with brown pigment, c. 1-5-1-7 /xm wide;<br />
ascogenous hyphae with swollen cells up to 5 ^tm wide. Excipulum reflexed but distinct, reddish<br />
brown but darker than <strong>the</strong> hypo<strong>the</strong>cium; hyphae radiating, branched and anastomosing,<br />
l-l-5(-2) /xm diam, hyaline (pigment confined to gel-matrix).<br />
Pycnidia <strong>of</strong>ten present but very inconspicuous, immersed, 35-40 /u.m diam, wall dark sordid<br />
green, K— , HNO3+<br />
red. Conidia (microconidia) ± cylindrical, c. 3-5-4-7x1 /xm.<br />
Chemistry: Sections <strong>of</strong> thallus and apo<strong>the</strong>cia K- , C-<br />
, PD-<br />
; no substances detected by t. I.e.<br />
Observations: M. turfosa is fairly constant in appearance, except that <strong>the</strong> thallus is less well<br />
developed when in boggy habitats. Microscopically <strong>the</strong>re is much variation between collections<br />
with regard to spore septation: some having mostly simple spores (e.g. Vezda Lich. Sel. 538),<br />
o<strong>the</strong>rs having numerous 1-septate and several 2- or 3-septate spores (e.g. Vezda Lich. Sel.<br />
1135). In <strong>the</strong> type <strong>of</strong> Biatora turfosa <strong>the</strong> spores are mostly simple but a few with a single septum<br />
were found. In <strong>the</strong> type collections <strong>of</strong> Biatora turfosa* verrucula most spores are 1-septate.
LICHEN GENUS MICAREA IN EUROPE 195<br />
With <strong>the</strong> combination <strong>of</strong> blackish thallus and black, convex to ± globose apo<strong>the</strong>cia, M.<br />
turfosa is easily confused in <strong>the</strong> field with peat inhabiting forms <strong>of</strong> M. melaena, but <strong>the</strong> latter<br />
differs microscopically in having dark purple and/or green pigments in <strong>the</strong> hypo<strong>the</strong>cium and<br />
thinner paraphyses. M. botryoides has similar apo<strong>the</strong>cial pigmentation to M. turfosa, but its<br />
hypo<strong>the</strong>cium is darker with nearly all <strong>the</strong> hyphae densely coated with pigment; in addition, M.<br />
botryoides has less numerous and (when hyaline) thinner paraphyses, a shallower hymenium,<br />
smaller spores, and its apo<strong>the</strong>cia are always accompanied by numerous, stalked, black pycnidia.<br />
M. melaenida is unlikely to be mistaken for M. turfosa because <strong>of</strong> its whitish thallus and habitat<br />
<strong>of</strong> fine-grained, mineral soils; it can be fur<strong>the</strong>r distinguished by its purple-brown epi<strong>the</strong>cium,<br />
excipulum, and (usually) upper hypo<strong>the</strong>cium. The little known M. osloensis differs from M.<br />
turfosa in several features but is most easily distinguished by its much smaller, non-septate<br />
spores.<br />
Habitat and distribution: M. turfosa seems to have an arctic-alpine distribution. In Britain it is<br />
known from summits in <strong>the</strong> Cairngorm, Grampian, and Breadalbane mountains <strong>of</strong> Scotland,<br />
where it grows on exposed turf (mostly over dead bryophytes) at altitudes <strong>of</strong> about 860-1245 m.<br />
In <strong>the</strong> Sudeten and Carpathian mountains <strong>of</strong> Czechoslovakia it is reported at altitudes <strong>of</strong> 1400 m<br />
and 1870 m respectively. It seems to be able to occur at lower altitudes in Sphagnum-bogs in<br />
Scandinavia, but no altitude data is given with any <strong>of</strong> <strong>the</strong> specimens seen. I have not seen any<br />
material from <strong>the</strong> Alps but it is likely to occur <strong>the</strong>re.<br />
On <strong>the</strong> Cairngorm plateau M. turfosa occurs amongst Juncus trifidus. Associated lichens on<br />
<strong>the</strong> <strong>British</strong> ga<strong>the</strong>rings include Cladonia spp., Lecidea caesioatra, and Lepraria neglecta.<br />
Map 27 Micarea turfosa • 1950 onwards O Before 1950
196 BRIAN JOHN COPPINS<br />
Outside Europe M. turfosa is little known, but I have seen material from Greenland and<br />
nor<strong>the</strong>rn Alaska.<br />
Exsiccata: Flotow Lich. 130 (VER). Korber Lich. Sel. 12 (L). Magnusson Lich. Sel. Scand. 282 (GZU).<br />
Malme Lich. Suec. 865 (WIS). Norrlin & Nyl. Herb. Lich. Fenn. 321 (BM, M). Vezda Lich. Sel. 538 (BM),<br />
1135 (BM).<br />
Excluded taxa<br />
Bacidia beckhausii Korber, Parerga lich.: 134 (1860). - Micarea beckhausii (Korber) Vezda in Poelt &<br />
Vezda, Bestimmungsschl. europ. Flechten, Ergdnzungsheft I: 162 (1977). Type: Germany, 'West-<br />
phalen', Beckham (L 910, 137 1363 - lectotype!).<br />
Bacidia miniuscula Anzi, Cat. lich. Sondr. : 70 (I860). - Micarea miniuscula (Anzi) Vezda, in Vezda & V.<br />
Wirth in Folia geobot. phytotax., Praha 11: 100 (1976). Type: not seen.<br />
Bacidia beckhausii must be excluded from Micarea, mainly on account <strong>of</strong> its excipulum structure (see<br />
p. 189 and 198).<br />
Bacidia nitscltkeana var. perpusilloides Erichsen in Annals mycol. 41: 206 (1943). Type: West Germany,<br />
Schleswig-Holstein: Flensburg, Forst Clusries, near Wasserleben, on Picea twigs, 6x1923, C. F. E.<br />
Erichsen (HBG - lectotype!). Paratypes: Schleswig-Holstein: Flensburg, Jerrishoer Holz, 10 vi 1928,<br />
C. F. E. Erichsen (HBG!); Hamburg, Wohldorfer Wald, 12 vi 1905, C. F. E. Erichsen (HBG!);<br />
Hamburg, Volksdorfer Wald, 19 xii 1909, C. F. E. Erichsen (HBG!). All on young twigs <strong>of</strong> Picea.<br />
Thallus thin, green-grey. Apo<strong>the</strong>cia minute, ± pellucid, pale brown, immarginate, c. 0-1-0-15 mm diam.<br />
Hymenium, hyaline, c. 30-40 /Am tall. Hypo<strong>the</strong>cium dark brown, K-, or -I- greenish in part, c. 40-45 /im<br />
tall. Excipulum thin, c. 12-15 /xm wide, ± pseudoparenchymatous with cells c. 3-5-8x2-5-3 /Am (in K),<br />
hyahne or with greenish pigment in zone adjacent to hymenium. Asci clavate, numerous, 8-spored. Spores<br />
oblong-fusiform, straight or slightly curved, 3(-5)-septate, 16-24(-28)x3-4 /^m (Fig. 56A). Pycnidia <strong>of</strong>ten<br />
numerous, blackish, c. 50 /Am diam, wall greenish in K; conidia curved or sigmoid 12-30x0-8 /tm.<br />
Phycobiont cells c. 5-10 /im diam.<br />
This variety agrees in all respects, except host substratum, with Bacidia myrtillicola Erichsen in Mitt.<br />
Inst. allg. Bot. Hamb. 10: 414 (1939); type: Schleswig-Holstein: Lauenburg, 'im "Sachsenwald" bei<br />
Friedrichsruh an Vaccinium myrtillus am Rande des Reviers "Saupark".', 2 xi 1924, C. F. E. Erichsen<br />
(HBG - holotype!). B. myrtillicola belongs to <strong>the</strong> mainly foliicolous groups <strong>of</strong> Bacidia s. lat. that includes<br />
B. fuscatula (Miill. Arg.) Zahlbr., B. rhapidophylli (Rehm) Zahlbr., and B. vezdae Coppins & P. James.<br />
Catillaria melanobola f. frullaniae B. de Lesd., Rech. Lich. Dunkerque, Suppl. 1: 119 (1914). Type:<br />
France, Nord, near Burgues, on Frullania on Ulmus, M. B. de Lesdain, Zahlbr. Lich. Rar. Exs. 152 (BM<br />
-isotype!).<br />
This is an apparently undescribed (at specific rank) species <strong>of</strong> Arthonia s. lat. The apo<strong>the</strong>cia are very<br />
small, c. 0- 1 mm diam, convex-globose and black. In section <strong>the</strong> epi<strong>the</strong>cium and hypo<strong>the</strong>cium are brownish<br />
(K-l- olive), <strong>the</strong> asci clavate, c. 21 x 10 /tm at maturity, with a minute internal amyloid ring [as described for<br />
Bryostigma leucodontis (Poelt & Dobbeler, 1979) and Chrysothrix (Laundon, 1981), and noted by me in<br />
several Arthonia species, especially lichenicolous species and members <strong>of</strong> <strong>the</strong> subgenus Allarthonia],<br />
spores hyaline, 1-septate, 9-11x2-3-3 /im, and paraphyses scanty, thin, c. 0-6-1 /Am wide. This species is<br />
ra<strong>the</strong>r common in <strong>the</strong> <strong>British</strong> Isles where it is usually referred to as 'Arthonia cf. exilis\ It is found in a<br />
variety <strong>of</strong> habitats, for example bark <strong>of</strong> shaded trunks <strong>of</strong> Acer, Fraxinus and Ulmus, branches and twigs <strong>of</strong><br />
Corylus and Sambucus, over bryophytes (especially Frullania) on trunks <strong>of</strong> Quercus and Ulmus, tufts <strong>of</strong><br />
Armeria on sea-cliffs, and possibly also on sandstone rocks. When on bark <strong>the</strong> apo<strong>the</strong>cia may attain a larger<br />
size, to 0-2 mm diam. Corticolous material has recently been distributed as Vezda Lich. Sel. 1701. In E it is<br />
represented by material from <strong>the</strong> following <strong>British</strong> vice-counties: 1, 20, 29, 31 , 62, 64, 67, 78, 83, 88, 90, 98,<br />
99, 110, H18 and H20; and from Denmark and Germany (Schleswig-Holstein).<br />
There has been much confusion regarding <strong>the</strong> correct interpretation <strong>of</strong> Arthonia exilis (Florke) Anzi. I<br />
have recently had <strong>the</strong> opportunity <strong>of</strong> examining type material <strong>of</strong> this species (Florke Deutsch. Lich. 187<br />
(WRSL - lectotype!)) and a brief description <strong>of</strong> it is as follows:<br />
Thallus lignicolous (probably on worked timber; accompanied by a few granular-areolae <strong>of</strong> Candelariella<br />
vitellina), thin, grey-white. Phycobiont noX Trentepohlia, cells, 10-19 /Am diam. Apo<strong>the</strong>cia numerous, ±<br />
regularly dispersed, black, 0-1-0-2 mm diam. Epi<strong>the</strong>cium (including lower border with substratum)<br />
reddish brown, K-. Hymenium c. 30 /im tall, I -I- reddish, K/I-l- blue. ^5d clavate, with minute amyloid<br />
ring ("Bryostigma -type'), 22-30x14-15 /im, 8-spored. Spores hyahne, 1-septate, 8-12x2-5-3-5(^) /Am.<br />
Paraphyses ± coherent (even in K), ra<strong>the</strong>r stout, c. 1-5-2 /xm wide; apices ± swollen with brown
LICHEN GENUS MICAREA IN EUROPE 197<br />
Fig. 56 A, spores <strong>of</strong> Bacidia myrtillicola (HBG - lectotype <strong>of</strong> Bacidia nitschkeana var. perpusilloides).<br />
B-C, Catillaria bouteillei (H-NYL 1884 - holotype <strong>of</strong> Lecidea littorella); B, spores; C, conidiogenous<br />
cells and conidia. Scale = 10 /u,m.<br />
pigment-caps; closely adhering pigment sometimes continuing down to about mid-hymenium level.<br />
Hypo<strong>the</strong>cium hyaline.<br />
The status oiA. exilis s. str. as a <strong>British</strong> species requires confirmation.<br />
Catillaria rhodosphaeraTh . Fr. & HultinginTh. Fr. Lich. Scand. 2: 571 (1874). Type: Sweden, Dalsland,<br />
Haverud, 1870, /. Hulling (UPS - lectotype! ; UPS<br />
- isolectotype!).<br />
This species was provisionally referred to Micarea by Kilias (1980: 392), but in fact belongs to Biatora<br />
Fr., a name which requires conservation against Biatora Ach. (= Stenhammarella Hertel). In my opinion<br />
C. rhodosphaera represents a saxicolous form <strong>of</strong> Catillaria sphaeroides (Dickson) Schuler.<br />
Lecidea arceutina var. hypnaea Nyl. in Flora, Jena 51: 165 (1968). - Bacidia arceutina f . hypnaea (Nyl.) A.<br />
L. Sm. , Monogr. Brit. Lich. 2: 158 (1911). Type: Jersey, 1866, Larbalestier {W-]
198 BRIAN JOHN COPPINS<br />
arceutina (Ach.) Arnold. In his diagnosis Nylander gives <strong>the</strong> spore length as 45-70 /xm and on <strong>the</strong> holotype<br />
packet he wrote '45-20x1 yam'; my examination <strong>of</strong> <strong>the</strong> holotypefound <strong>the</strong> spores to be 38-51x1-5 /xm.<br />
Lecidea clavulifera Nyl. - see Psilolechia clavulifera.<br />
Lecidea demarginata Nyl. in Flora, Jena 61: 245 (1878). Type: Finland, Tavastia australis, Evo, 'supra<br />
saxum in sylva juniori, loco olim deusto', 1873,7. P. Norrlin, Norrlin & Nyl. Herb. Lich. Fenn. 179 (Hlectotype!;<br />
isolectotypes (without exsiccate label): H-NYL 20162! & 20163!).<br />
In <strong>the</strong> protologue Nylander cited material <strong>of</strong> Norrlin's from Finland and <strong>of</strong> Larbalestier's from Ireland.<br />
The epi<strong>the</strong>t 'demarginata' was first used in 1875 on <strong>the</strong> label <strong>of</strong> Herb. Lich. Fenn. 179, although without a<br />
diagnosis. An example <strong>of</strong> this exsiccate is here selected as lectotype, and is a specimen <strong>of</strong> Lecidea erratica<br />
Korber. The cited Larbalestier material is represented in BM and H and belongs to M. lutulata.<br />
Lecidea denigrata var. bacidiella Vainio in Medd. Soc. Fauna Fl. fenn. 10: 28 (1883). - Micarea bacidiella<br />
(Vainio) Vezda, in Vezda &. V. Wirth in Folia geobot. phytotax., Praha 11: 100 (1976). Type: Finland,<br />
Lapponia kemensis, Sodankyla, Pyhatunturi, Kannolla, on hgnum, 1878, E. A. Vainio (TUR-VAINIO<br />
22505 -holotype!).<br />
As stated by Vainio (1934: 462) this is a synonym <strong>of</strong> Bacidia miniuscula Anzi (i.e. B. beckhausii Korber).<br />
The following were notes made from <strong>the</strong> holotype <strong>of</strong> var. bacidiella:<br />
Thallus lignicolous, entirely endoxylic. Phycobiont cells c. 8-14 ^tm diam. Apo<strong>the</strong>cia thinly marginate<br />
and ± plane when young, soon becoming convex and immarginate, black, c. 0-2-0-3 mm diam. Hymenium<br />
c. 35 /xm tall, with olivaceous vertical streaks, K+ violet, C+ violet (not red). Asci clavate, 30-35x10-12<br />
/im, 8-spored. Spores rod-shaped or slightly curved, (l-)3(-7)-septate, 17-26x1-7-2 /nm. Paraphyses<br />
simple or sparingly branched, 1-1-5 /xm wide; apices somewhat incrassate, to 2 /xm wide. Hypo<strong>the</strong>cium<br />
hyaline. Excipulum dilute olivaceous (K+ violet), <strong>of</strong> radiating, branched hyphae c. 1-5-2 /xm wide<br />
embedded in a dense gel; hyphae becoming more distinct in K but not separating. Pycnidia immersed in <strong>the</strong><br />
substratum, black, c. 50-100 /xm diam; walls olivaceous, K-l- violet. Conidia simple, hyaline, oblong-<br />
ellipsoid, <strong>of</strong>ten wider at proximal end, 2-8-3-5 x 1(-1 -4) /xm.<br />
B. beckhausii usually occurs on bark, although I have seen a few additional lignicolous specimens from<br />
Scotland. Micarea bacidiella sensu Vezda & Wirth {loc. cit. ) is not B. beckhausii but <strong>the</strong> superficially similar<br />
Micarea globulosella (see p. 135).<br />
Lecidea erysiboides Nyl. in Not. Sdllsk. Fauna Fl. fenn. 4: 232 (1859). - Catillaria erysiboides (Nyl.) Th. Fr.<br />
Lich. Scand. 2: 572 (1874). Type: Finland, Nylandia, Helsingfors [Helsinki], 'Gumtackt' on rotting<br />
decorticate pine trunk, 1858, W. Nylander (H - lectotype!; H-NYL p.m. 4514- isolectotype!).<br />
This name has been misapplied by many lichenologists and <strong>the</strong> majority <strong>of</strong> specimens seen belong to<br />
Micarea prasina, or o<strong>the</strong>r species such as Catillaria sphaeroides, Lecania cyrtella, and Micarea denigrata.<br />
The type material <strong>of</strong> L. erysiboides is not a Micarea but a species <strong>of</strong> Catillaria s. lat. , although I am<br />
uncertain <strong>of</strong> its affinities. It has small, plane to convex, reddish brown (testaceous) apo<strong>the</strong>cia that are<br />
marginate when young, <strong>the</strong> excipulum is composed <strong>of</strong> much-branched, radiating hyphae which are distinct<br />
in K but still tightly bound by <strong>the</strong> gel matrix, and <strong>the</strong> spores are ovoid and <strong>of</strong>ten constricted at <strong>the</strong> septum,<br />
1-septate with <strong>the</strong> upper cell usually enlarged and ± globose, 8-9-5x3-5 /xm. C. erysiboides has not been<br />
correctly reported from <strong>the</strong> <strong>British</strong> Isles but it should be looked for, especially in <strong>the</strong> eastern Scottish<br />
highlands.<br />
Additional specimens <strong>of</strong> C. erysiboides examined: NORWAY. Hordland: Granvin, on Betula lignum,<br />
1904, Havaas, Lich. Exs. Norv. 292 (BG). FINLAND. Nylandia: Helsinki, 1859, W. Nylander (H-NYL<br />
21650), and 1861 (BM). ITALY: Trentino ('Sudtirol'): Paneveggio, on top <strong>of</strong> cut stump <strong>of</strong> Picea, 2 ix 1883,<br />
Arnold, Lich. Exs. 1002 (BM ex K). USSR. 'Lapponia orientalis', 1863, A^. /. Fellman, Lich. Arct. 156<br />
(BMexK).<br />
Lecidea fuliginea Ach., Syn. Lich.: 35 (1814). - Micarea fuliginea (Ach.) Fr., Syst. orb.: 257 (1825). -<br />
Micarea fuliginea (Ach.) Fr., Stirp. agrifemsion. : 37 (1825); comb, inval. (Arts 34.1, 43).<br />
I have not seen <strong>the</strong> type <strong>of</strong> this name but it is probably a synonym <strong>of</strong> Lecidea icmalea Ach. A later<br />
synonym <strong>of</strong> <strong>the</strong> latter is Pannularia perfurfurea Nyl. which is <strong>the</strong> type <strong>of</strong> <strong>the</strong> genus Placynthiella Gyelnik.<br />
Lecidea gelatinosa Florke in Magazin Ges. naturf. Fr. Berl. 3: 201 (1809). - Micarea gelatinosa (Florke)<br />
Brodo in Bryologist 70: 216 (1967).<br />
I have not seen type material <strong>of</strong> this species, but if its general interpretation is correct <strong>the</strong>n it is not a<br />
Micarea. In fact it belongs in <strong>the</strong> Lecidea granulosa group, which includes L. aeruginosa Borrer, L.<br />
aene<strong>of</strong>usca Florke ex Flotow, L. granulosa (H<strong>of</strong>fm.) Ach. and L. viridescens (Schrader) Ach. These<br />
species are not congeneric with <strong>the</strong> type species <strong>of</strong> Lecidea Ach. (L. fuscoatra (L.) Ach.) and <strong>the</strong>y await <strong>the</strong><br />
formal transference to a genus in <strong>the</strong> Trapeliaceae . Fur<strong>the</strong>r detailed studies are needed to see if <strong>the</strong>y can be 1
LICHEN GENUS MICAREA IN EUROPE 199<br />
placed in Trapelia Choisy or Trapeliopsis Hertel & G. Schneider, or if a new genus is required to<br />
accommodate <strong>the</strong>m.<br />
Lecidea littorella Nyl. in Flora, Jena 60: 229 (1877). - Catillaria littorella (Nyl.) Zahlbr., Cat. lich. univ. 4:<br />
56 (1926). Type: Ireland, West Galway, 'Hibernia occidentalis, Bord du Lough Inagh, mais extremement<br />
rare', 1876, C. Larbalestier (Yi-^Yl. 18884- holotype!, isotypes: BM!, BM ex K!).<br />
The following notes were made from <strong>the</strong> holotype: Thallus saxicolous, not delimited but forming small<br />
patches amongst Hymenelia lacustris and Porina chlorotica, whitish, or dull ochraceous in part (? due to<br />
age), rimose, matt. Apo<strong>the</strong>cia numerous, weakly marginate, plane to slightly convex, pallid to dull<br />
orange-red, 0-1-0-3 mm diam; margin not exceeding <strong>the</strong> level <strong>of</strong> <strong>the</strong> disc, very thin, c. 0-1-0-3 mm diam;<br />
margin not exceeding <strong>the</strong> level <strong>of</strong> <strong>the</strong> disc, very thin, c. 0-01-0-02 mm wide, whitish with faintly pruinose<br />
appearance. Hymenium 40-47 jxxn tall, hyaline. Asci cylindrical-clavate, ' Lecanora-type\ 8-spored.<br />
Spores ovoid to ovoid-oblong, <strong>of</strong>ten 'slipper-shaped', 1-septate and <strong>of</strong>ten constricted at <strong>the</strong> septum,<br />
9-5-14(-16)x4-6(-7) ^tm (Fig. 56B). Paraphyses numerous, thin, 0-8-1 ixm wide, simple or sparingly<br />
branched below, becoming wider (to 1-7 ixm) and more frequently branched above. Hypo<strong>the</strong>cium hyaline.<br />
Excipulum hyaline or dilute yellow-straw, inspersed with minute crystals in water mounts but ± clearing in<br />
K, c. 20 /am wide laterally, slightly widening below to 28 )u,m, minutely paraplechtenchymatous with cells in<br />
<strong>the</strong> lower part measuring 4—6x2-5^ /u,m. Pycnidia frequent, at first immersed, later becoming emergent,<br />
white (translucent when wet), c. 80-100 fxm diam. Conidia (Fig. 56C) simple, ovoid to pyriform,<br />
2-8xl-5-l-7)Ltm.<br />
L. littorella was provisionally referred to Micarea by Kilias (1981: 392). However, my subsequent<br />
examination <strong>of</strong> <strong>the</strong> holotype proved it to be a saxicolous form <strong>of</strong> <strong>the</strong> normally foliicolous Catillaria<br />
bouteillei (Desm.) Zahlbr.; previous reports <strong>of</strong> this species on rock are given by Degelius (1944) and<br />
Santesson (1952). C. bouteillei is apparently widely distributed in Ireland, especially on <strong>the</strong> leaves oiBuxus<br />
(Knowles, 1929; Scannell, 1978). An additional <strong>British</strong> saxicolous specimen has been located: V.C.5,<br />
South Somerset, Broomfield, Ruborough Camp, 1914, W. Watson (BM ex K).<br />
Nylander's choice <strong>of</strong> <strong>the</strong> epi<strong>the</strong>t 'littorella' is ra<strong>the</strong>r misleading because <strong>the</strong> holotype was not collected on<br />
<strong>the</strong> sea-shore, but on <strong>the</strong> shores <strong>of</strong> an inland, freshwater lake some 10 km from <strong>the</strong> sea. <strong>British</strong> authors (e.g.<br />
James, 1970; Fletcher, 1975) have mistakenly applied <strong>the</strong> name Catillaria littorella to a species that grows in<br />
crevices in siliceous rocks near <strong>the</strong> sea-shore; this species probably belongs to <strong>the</strong> perplexing Lecania<br />
ery5/fte complex.<br />
Lecidea milliaria a. [var.] terrestrisFr. Lich. Eur. 342 (1831). Type: not designated.<br />
This name is <strong>of</strong> unlikely appHcation in M/carea. Fries cited his L/c/i. Suec. no. 213, <strong>the</strong> example <strong>of</strong> which<br />
in UPS is a member <strong>of</strong> <strong>the</strong> Lecidea limosa group.<br />
Lecidea ocelliformis Nyl. in Flora, Jena 48: 145 (1865). - Bilimbia ocelliformis (Nyl.) Branth. & Rostr. in<br />
Bot. Tidskr. 3: 226 (1869). - Lecidea atroviridis f. ocelliformis (Nyl.) Hedl. in Bih. K. svenska<br />
VetenskAkad Handl. Ill, 18 (3): 64 (1892). - Catillaria prasina f. ocelliformis (Nyl.) Erichsen in Annls.<br />
mycol. 41: 205 (1943). Type: Finland, Tavastia australis, Hollola, ad corticem Sorbi, 30 vi 1863, /. P.<br />
Norrlin 210 (H-NYL 20607! - labelled Tsotype' by M. Inoue in 1979).<br />
This is not a Micarea. If <strong>the</strong> types <strong>of</strong> L. atroviridis (Arnold) Th. Fr. and L. ocelliformis are conspecific, as<br />
placed by Hedlund (op. cit.) and Vainio (1934: 218), <strong>the</strong>n <strong>the</strong> latter epi<strong>the</strong>t has priority.<br />
Lecidea pauxilla Krempelh. in Verh. zool.-bot. Ges. Wien 26: 455 (1876). Type: New Zealand, on bark<br />
[? Podocarpus], 18 -, C. Knight (M - lectotype!; M - isolectotype!).<br />
L. pauxilla was given as a synonym <strong>of</strong> Catillaria syno<strong>the</strong>a (i.e. Micarea denigrata) by Zahlbruckner (Cat.<br />
lich. univ. 4: 78, 1926). However, <strong>the</strong> type material belongs to Cliostomum griffithii (Sm.) Coppins.<br />
Lecidea recondita Erichsen in Annls mycol. 42: 25 (1944). Type: Germany, Schleswig-Holstein: 'Apen-<br />
rade, an Blocken am Grunde einer Erosionsschluct im Geholz Jiirgensgaard' [now Denmark, Jylland,<br />
near Abenra], 28 vii 1932, C. F. E. Erichsen (HBG - holotype!).<br />
In <strong>the</strong> protologue Erichsen compares this with Lecidea sylvicola (= Micarea sylvicold). My examinations<br />
<strong>of</strong> <strong>the</strong> holotype proved it not to be a Micarea but referable to Catillaria chalybeia (Borrer) Massal.; see<br />
Kilias (1980: 448).<br />
Lecidea sabuletorum f. simp/icior Nyl., Lich. Scand.: 205 (1861). - Bilimbia trisepta f. simplicior (Nyl.)<br />
Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 258 (1922). Type: not designated.<br />
Nylander (loc. cit.) cited <strong>the</strong> following material 'supra muscos ad Helsingforse [ipse] et in Sueciae<br />
montibus [Thedenius] atque supra terram in Lapponia'.<br />
According to Fries (1874: 523) <strong>the</strong> 'Helsingforse' specimen is Lecidea verrucula (= Micarea turfosa);<br />
according to Vainio (1922: 140) <strong>the</strong> Swedish specimen from Funnesdalsberget in Harjedalen collected by
200 BRIAN JOHN COPPINS<br />
K. F. Thedenius is Lecidea dufourei (= Catillaria contristans (Nyl.) Zahlbr.). I am uncertain as to <strong>the</strong><br />
identity <strong>of</strong> <strong>the</strong> specimen(s) referred to by Nylander as 'in Lapponia'. This may be based on <strong>the</strong> specimen<br />
cited by Vainio (1922: 140, 258) as 'supra Grimmias in Kipina Lapponiae a G. SeHn coUecta', which Vainio<br />
refers to Bilimbia trisepta [sensu Vainio = Micarea peliocarpa] f. simplicior. However, material in<br />
Nylander's herbarium (H-NYL 18841) labelled 'Lapponia, G. Selin 1861' is Catillaria contristans.<br />
Fur<strong>the</strong>rmore, ano<strong>the</strong>r specimen <strong>of</strong> 'Lecidea sabuletorum f. simplicior' collected from Iceland by Isaac<br />
Carroll in 1861 (H-NYL 18862) is also C. contristans. It would appear that Nylander's concept <strong>of</strong><br />
'simplicior' is best fitted by C. contristans and should be typified on <strong>the</strong> material collected by Thedenius or<br />
Selin.<br />
Lichen niger Hudson, Fl. angl., ed. 2, 2: 524(1778). -Collemanigrum{Hudson)Y{<strong>of</strong>fm. , Deutschl. Fl.: 103<br />
(1796). - Micarea nigra (Hudson) Fr., Syst. orb.: 257 (1825). - Micarea nigra (Hudson) Fr., Stirp. agri<br />
femsion.: 37 (1825); comb, inval. (Arts 34.1, 43). - Placynthium nigrum (Hudson) Gray, Nat. Arr. Br.<br />
PI. 1:395(1821).<br />
This is <strong>the</strong> type species <strong>of</strong> <strong>the</strong> genus Placynthium Gray.<br />
Lichen viridescens Schrader, in Gmelin Syst. nat. 2 (2): 1361 (1792). - Lecidea viridescens (Schrader) Ach.<br />
Meth. Lich. : 62 (1803). - Micarea viridescens (Schrader) Brodo in Bryologist 70: 216 (1967).<br />
I have not seen <strong>the</strong> type <strong>of</strong> this name but if its normal interpretation is correct <strong>the</strong>n <strong>the</strong> species belongs to<br />
<strong>the</strong> Lecidea granulosa group; see notes above under Lecidea gelatinosa.<br />
Micarea chrysophthalmaF . James in Lichenologist 5: 131 (1971). - Chrysothrix chrysophthalma (P. James)<br />
P. James & Laundon, in Laundon in Lichenologist 13: 104 (1981).<br />
This is a species <strong>of</strong> Chrysothrix and is discussed in detail by Laundon (1981).<br />
Micarea coccinea Fr., Syst. orb. :<br />
257 (1825); nom. nudum (Art. 32.1).<br />
I do not know <strong>the</strong> identity <strong>of</strong> this species, and I have not seen this name in any o<strong>the</strong>r publication. On<br />
morphological grounds it is unlikely that Fries intended a new combination based on Lichen coccineus<br />
Dickson [= Haematomma ochroleucum (Necker) Laundon] or Lecidea coccinea Schw. [= Lecidea russula<br />
Ach.].<br />
Micarea cyanescens Poelt & Dobbeler in Bot. Jb 96: 339 (1975). Holotype: Germany, Bayern, Altbayerische<br />
Alpen, Chiemgauer Berge, 900-1000 m on Campylium halleri on north-facing calcareous rocks, 29<br />
ix 1974, /. Poelt (GZU, not seen). Paratypes: Austria, Tirol, Lechtaler Alpen, 'unteres Medriol-Tal<br />
uber Zams bei Landeck, an Dolomitblocken unter Griinerlen', 1600-1700 m, on Campylium halleri,<br />
1969, J. Poelt (hb Poelt 13295!); Austria, Steiermark, 'uber Kalk, Rote Wand bei Mixnitz, c. 1470 m,<br />
unweit des Gipfels', 1974, J. Poelt (GZU!).<br />
This is a very curious species; its hymenium components are embedded in a dense gel which does not<br />
disperse in 50% <strong>of</strong> KOH. It is not a Micarea and a new genus is probably required to accommodate it.<br />
Micarea hylocomii Poelt & Dobbler in Bot. Jb. 96: 341 (1975). Type: Austria, Tirol, Samnaun Gruppe, on<br />
<strong>the</strong> way from Serfaus to Madatschen, c. 1500 m, on Hylocomium splendens, 15 ix 1972, J. Poelt (GZU -<br />
holotype!).<br />
I endorse <strong>the</strong> describing authors' view that <strong>the</strong> position <strong>of</strong> this species in Micarea is doubtful, and I<br />
beheve a new genus should be erected to accommodate it.<br />
Micarea leprosa P. James in Lichenologist 5: 133 (1971). - Vezdaea leprosa (P. James) Vezda, in Poelt &<br />
Dobbeler in Lichenologist 9: 170 (1977).<br />
This is a species <strong>of</strong> Vezdaea (Poelt & Dobbeler, 1975). A large collection <strong>of</strong> it from Scotland has recently<br />
been distributed as Hertel Lecid. Exs. 60.<br />
Micarea minima Poelt & Dobbeler in Bot. Jb. 96: 342 (1975). Holotype: Germany, Rheinland-Pfalz,<br />
'Boschung eines Waldweges bei Battweiler, Kreis Zweibriicken, c. 400 m', on Polytrichum commune<br />
and P. formosum, 1974, J. Poelt (GZU, not seen). Paratype: Austria, Karnten, Nockgruppe, Afritzer<br />
Berge, 'Bergwald und saure Weiden iiber Verditz, nordlich Villach', c. 1300 m, on Polytrichum<br />
formosum, 1974,/. Poe/f (hb Poelt 13294!).<br />
I was unable to find any apo<strong>the</strong>cia in <strong>the</strong> examined paratype and so I cannot confidently exclude this<br />
species from Micarea. Fur<strong>the</strong>r studies are required to establish its generic disposition.<br />
Patellaria nigrataMuW. Arg. in Flora, Jena7l: 540(18SS).-Bacidianigrata{Mu\\. Arg.) Zahlbr., Car. lich.<br />
univ. 4: 129 (1926). - Micarea nigrata (Miill. Arg.) Kalb, Lichenes Neotropici Fasc. 1: 8 (1981); comb,<br />
inval. (Art. 33.2). Type: Brazil, Apiahy, on argillaceous soil, 1883, Puiggari (G!).<br />
This species closely resembles a Micarea and even appears to have 'micareoid' algae. However, its
LICHEN GENUS MICAREA IN EUROPE 201<br />
excipular hyphae are conglutinated, broad (c. 2-2-5 /u,m in K), and pachydermatous. It is closely allied to<br />
Bacidia subrudecta (Vainio) Zahlbr. and may have affinities with Byssoloma, but I think it is best retained<br />
in Bacidia s. lat. pending fur<strong>the</strong>r study. The material distributed by Kalb {Lich. Neotrop. Exs. ^2) as<br />
'Micarea nigrata' is M. lignaria var. lignaria.<br />
Psilolechia clavulifera (Nyl.) Coppins, comb. nov. - Lecidea clavulifera Nyl. in Flora, Jena 52: 294 (1869). -<br />
Micarea clavulifera (Nyl.) Coppins & P. James, in D. Hawksw. , P. James & Coppins in Lichenologist 12:<br />
107 (1980). Type: Finland, Lapponia kemensis, Muonionska [Muonio], Keimioniemi, ad radicem<br />
abietis, 1867, J. P. Norrlin 627 (H - lectotype!; isolectotypes: H!, H-NYL 20855!).<br />
Lecidea clavulifera f. subviridicans Nyl. in Flora, Jena 60: 463 (1877). Type: Ireland, West Galway,<br />
Connemara, Kylemore, in a cave on <strong>the</strong> NW side <strong>of</strong> Doughraugh mountain, 1876, C. Larbalestier (BMlectotype!<br />
Isolectotypes: BM ex K!; also distributed as Larbal. Lich. Herb. 29: BM!, H!).<br />
Thallus effuse, whitish or greenish white, <strong>of</strong> dispersed to coalescing, irregular granular-areolae c.<br />
Fig. 57 Psilolechia clavulifera (Coppins 3614, E). A, spores. B, paraphyses. C, phycobiont cells (cell<br />
contents and mycobiont hyphae omitted). Scale = 10 /xm.
202 BRIAN JOHN COPPINS<br />
0-1-0-2 mm diam. Areolae <strong>of</strong>ten disintegrating to form a scurfy-granular crust. Phycobiont not micareoid;<br />
cells in clusters and tightly bound by short-celled hyphae, but haustoria not observed; clusters interconnected<br />
by filamentous hyphae c. 1-5-2 ju.m wide. Phycobiont cells irregularly globose, broadly ellipsoid or<br />
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).<br />
Apo<strong>the</strong>cia convex-hemispherical and immarginate from <strong>the</strong> beginning, sometimes becoming subglo-<br />
bose, more rarely tuberculate, grey-black with bluish tinge, but shade forms sometimes whitish, blue-grey<br />
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<br />
radiating hyphae. Hymenium 28-35 /xm tall, dilute straw (shade forms), or dilute greenish or aeruginose<br />
(K-l- green intensifying) especially in <strong>the</strong> upper part. Asci cylindrical-clavate, 25-30x5-7 /im, 8-spored.<br />
Spores ovoid, oblong-ovoid or dacryoid, simple, 3-6x(l-)l-2-l-7(-2) /xm (Fig. 57A). Paraphyses (Fig.<br />
57B) numerous, usually branched, sometimes anastomosing, distinctly septate and appearing ± articu-<br />
lated, stout, 1-3-2 /Ltm wide; apices sometimes ± clavate and up to 3 jxm wide, walls not pigmented but<br />
<strong>of</strong>ten surrounded by deeply pigmented gel matrix. Hypo<strong>the</strong>cium c. 50-70 /tim tall, hyaline, or dilute<br />
greenish or aeruginose (K— , HNO3-I- red) but <strong>the</strong>n never darker than <strong>the</strong> hymenium. Excipulum<br />
indistinct, <strong>of</strong> radiating hyphae that protrude as loose, hyaline hyphae c. 1-5-2 /xm wide and up to 50 /x,m<br />
long.<br />
Pycnidia not found.<br />
Chemistry: All parts K- , KC- , C- , PD- ; no substances detected by t.l.c.<br />
Preliminary studies <strong>of</strong> Lecidea clavulifera by Mr P. W. James and myself led us to transfer it to Micarea.<br />
However, critical studies have given me second thoughts on this placement. The distinct white rim <strong>of</strong><br />
protruding excipular hyphae (superficially like those <strong>of</strong> Byssoloma spp.), stout and distinctly septate<br />
paraphyses, and <strong>the</strong> unusual phycobiont are all uncharacteristic <strong>of</strong> a Micarea. The consideration <strong>of</strong><br />
Psilolechia Massal. as an alternative genus for L. clavulifera was prompted by observations <strong>of</strong> a<br />
lichenicolous member <strong>of</strong> <strong>the</strong> CaHciales, Microcalicium arenarium (Hampe ex Massal.) Tibell; M. arenarium<br />
is usually found as a parasite <strong>of</strong> Psilolechia lucida (Ach.) M. Choisy (see Tibell, 1978), but at three<br />
localities in Scotland I have found it on L. clavulifera. At <strong>the</strong> Berwickshire locality <strong>the</strong> host and parasite<br />
occurred in abundance on roots, stones, and soil <strong>of</strong> two up-ended trees (Fraxinus); P. lucida was also<br />
present in quantity but was not parasitized by M. arenarium.<br />
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 ±<br />
identical anatomical features, e.g. nature <strong>of</strong> paraphyses, size and shape <strong>of</strong> asci and spores, and excipulum<br />
with numerous protruding hyphae. P. lucida differs from P. clavulifera in <strong>the</strong> yellow-green colour <strong>of</strong> its<br />
leprose-granular thallus and apo<strong>the</strong>cia (due to presence <strong>of</strong> pulvinic acid derivatives), and a different<br />
phycobiont with ± globose cells 5-14 /xm diam. The thallus hyphae <strong>of</strong> both species are identical in<br />
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<br />
(in Poelt & Vezda, 1981). The phycobiont <strong>of</strong> P. clavulifera is very unusual; it is reminiscent <strong>of</strong> <strong>the</strong><br />
Stichococcus phycobiont <strong>of</strong> species such as Chaeno<strong>the</strong>ca stemonea and Coniocybe furfuracea, but its cells<br />
are much larger.<br />
P. clavulifera occurs in communities <strong>of</strong> <strong>the</strong> Micareetum sylvicolae, on roots, stones, and consolidated soil<br />
<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<br />
it is also known from Cumbria, south Wales and western Ireland (Map 28). From outside Britain I have<br />
seen material <strong>of</strong> it from Finland, Germany, and Czechoslovakia (see below).<br />
Additional specimens <strong>of</strong> P. clavulifera examined: <strong>British</strong> Isles. Brecon (V.C.42). 22/94: Upper<br />
Dyfnant Valley, 1982, Woods (hb Woods). Cumberland (V.C.70). 35/54: Baron Wood, by River Eden,<br />
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><br />
Whiteadder Water, exposed roots <strong>of</strong> up-ended Fraxinus, 1981, Coppins 8890 (E - Microcalicium<br />
arenarium folder). West Perth (V.C.87). 27/40: Aberfoyle, <strong>the</strong> Trossachs, on stone amongst roots <strong>of</strong><br />
up-ended tree, 1978, Coppins3661 (E). Mid Perth (V.C. 88). 27/55: Black Wood <strong>of</strong> Rannoch, stone in bank<br />
by track, 1976, Coppins 4654 (E); 27/81: Crieff, Drummond Wood, roots and sandstone <strong>of</strong> up-ended tree,<br />
partly parasited by Microcalicium arenarium, 1978, Coppins 3164, 3625 (E). Angus (V.C. 90). 37/33:<br />
Sidlaw Hills, Auchterhouse Hill, 396 m, underside <strong>of</strong> boulder, 1975, Coppins 846 (E - Microcalicium<br />
arenarium folder). South Ebudes (V.C. 102). 16/49: Colonsay, Coille Mhor, roots and soil <strong>of</strong> up-ended<br />
Betula, 1981, Coppins 8878 (E). Mid Ebudes (V.C. 103). Mull: 17/32: Bunessan, Ardfenaig Woods, 1970,<br />
James (BM); 17/54: Salen, 1968, James (BM); 17/55; Aros House, roots <strong>of</strong> up-ended tree, 1968, James<br />
(BM). West Galway (V.C.H.16). 84/72 (L/65): Connemara, near Clifden, 1878, Larbalestier (BM).<br />
Finland. Satakunta: Siikainen, Vuorijarvi, Vaasaneva, sandy soil amongst roots <strong>of</strong> Picea, 1936, Laurila<br />
(GZU, H). W. Germany. Baden-Wiirttemberg: Heidelberg, Konigstuhle, on sandstone, 1883, Zwackh<br />
(H-NYL p.m. 4229). Czechoslovakia. Slovakia, Nizke Tatry, Liptovska Teplicka, Dzurova, 18 -, Lojka<br />
4289 (BM).
LICHEN GENUS MICAREA IN EUROPE 203<br />
Map 28 Psilolechia clavulifera # 1950 onwards O Before 1950<br />
Sporacestra Massal. in Atti R. 1st. veneto Sci. Ill, 5: 264 (1860). Type species: Biatora prasina Tuck. &<br />
Mont, in Mont. (1857), non (Fr.) Trevisan (1856).<br />
Sporacestra was included as a synonym <strong>of</strong> Micarea by Vezda & Wirth (1976: 99) who confused <strong>the</strong> two<br />
quite separate applications <strong>of</strong> <strong>the</strong> name 'Biatora prasina' . The name Biatora prasina Tuck. & Mont, was<br />
introduced for a corticolous species from Venezuela with long acicular spores, and no reference was made<br />
to <strong>the</strong> 'prasina' <strong>of</strong> Fries. Similarly, Massalongo made no reference to Fries' name and his generic<br />
description (e.g. '.<br />
. . sporidii capillari aghiformi, lineari allungati . . .') clearly refers to <strong>the</strong> stated type<br />
species, ' Biatora prasina Mont., Tuck.'.<br />
The Venezuelan species was transferred to Bacidia de Not. by Zahlbruckner, but his use <strong>of</strong> <strong>the</strong> epi<strong>the</strong>t<br />
'prasina' is contrary to <strong>the</strong> present Code. From <strong>the</strong> descriptions <strong>of</strong> this species I believe it should be<br />
retained in Bacidia, pending fur<strong>the</strong>r study. The nomenclature and required new combination for <strong>the</strong><br />
species is as follows:<br />
Bacidia prasinata (Tuck.) Coppins, comb. nov. - Biatora prasinata Tuck, in Syn. N. Am. Lich 2: 41 (1888);<br />
nom. nov. - Bacidia prasina Tuck & Mont, in Mont, in Annls Sci. nat. IV, 8: 296 (1857), non (Fr.) Trevisan<br />
(1856). - Bacidia prasina Zahlbr., Cat. lich. univ. 4: 253 (1926); nom. illeg. (Art. 11). Type: Venezuela, on<br />
bark, Fendler (not seen).<br />
Index to exsiccatae<br />
I have attempted to examine at least one example <strong>of</strong> all European and North American material<br />
referable to Micarea and distributed in recognized exsiccatae (Lynge 1915-22, 1939; Sayre,<br />
1969). My preliminary list was compiled from Lynge {op. cit.), available schedae, major floras
204 BRIAN JOHN COPPINS<br />
(e.g. Fries, 1874; Korber, 1855, 1859-65; Smith, 1926; Vainio, 1922, 1934) and numerous o<strong>the</strong>r<br />
papers dealing with species here included in Micarea. Most examples gleaned from <strong>the</strong>se sources<br />
have been seen; <strong>the</strong> few not seen are listed after <strong>the</strong> main index. Herbarium locations for<br />
examined examples containing Micarea species can be found in <strong>the</strong> relevant taxonomic<br />
accounts; locations <strong>of</strong> examples not containing Micarea species are given in <strong>the</strong> index. The<br />
names on <strong>the</strong> exsiccate labels are given (in round brackets following <strong>the</strong> number) where <strong>the</strong>y are<br />
taxonomically at variance with <strong>the</strong> species represented.<br />
In many cases, only one or two examples <strong>of</strong> a particular number have been seen by me, thus<br />
this index must be regarded as provisional. Some numbers listed as not containing a Micarea<br />
(e.g. Zwackh Lich. Exs. US) may well in fur<strong>the</strong>r examples (not seen by me) contain a Micarea.<br />
In addition, some Micarea species may have been misidentified and distributed under names not<br />
connected with Micarea, and consequently overlooked in <strong>the</strong> course <strong>of</strong> this study.<br />
It should be noted that many numbers in <strong>the</strong> exsiccatae <strong>of</strong> Harmand {Lich. Loth.), Mougeot<br />
& Nestler, and Schaerer are notoriously heterogeneous with regard to species compo-<br />
sition, substratum, and locality. Most numbers in o<strong>the</strong>r exsiccatae are reasonably uniform in<br />
<strong>the</strong>se respects but heterogenicity sometimes arises in cases where species <strong>of</strong> similar outward<br />
appearance occur toge<strong>the</strong>r in <strong>the</strong> field (e.g. M. bauschiana-M. lutulata-M. sylvicola-M.<br />
tuberculata, and M. denigrata-M. lignaria-M. peliocarpa). During <strong>the</strong> preparation <strong>of</strong> exsiccatae<br />
<strong>the</strong> need for <strong>the</strong> careful identification <strong>of</strong> individual samples is paramount before <strong>the</strong>y are dis-<br />
tributed.<br />
Generic names are abbreviated as follows: Bacidia (B), Biatora (Bi.), Biatorina (Biat.),<br />
Bilimbia (Bit.), Catillaria (C), Lecanora (Lee), Lecidea (L.), Micarea (M.), Psilolechia {P.)<br />
and Scoliciosporum (5.).<br />
ANZI, Lich. Exs. Ital.: 256 M. denigrata; 259A (Bil. lignaria) M. melaena; 259B {Bil. hypnophila - given<br />
by Lynge as Bil. lignaria) B. sabuletorum (BM).<br />
ANZI, Lich. Lang.: 148 {Bil. syncomistd) M. lignaria, one example in BM with a little M. peliocarpa.<br />
ANZI, Lich. Sondr.: 170A M. peliocarpa; 170B M. melaena.<br />
ARNOLD, Lich. exs.: 120 M. bauschiana; 167A {Bil. lignaria ^saxigena Leighton) M. peliocarpa; 167B<br />
M. peliocarpa; 217 M. nitschkeana; 279, 280A-C M. prasina; 332A-C M. melaena; 348A, B M. lignaria;<br />
409A M. sylvicola; 409B (L. sylvicola) M. sylvicola, or M. lutulata; 503A-D M. nitschkeana; 548, 549 M.<br />
cinerea; 556[A], B (L. assimilata) M. crassipes; 626, 627 M. misella; 836 M. lithinella; 837 M. peliocarpa;<br />
1051 {Bil. ternaria) M. peliocarpa; 1057 M. tuberculata; 1121 M. crassipes; 1122 M. prasina; \A1\ M.<br />
elachista; 1472 M. prasina.<br />
ARNOLD, Lich. Mon.: 46 M. denigrata; 47 M. cinerea; 48 M. nitschkeana; 49 M. melaena; 115, 116 M.<br />
cinerea; 118 {Bil. trisepta) M. peliocarpa; 172 M. misella; 241 M. misella; 243 M. prasina; 244 {Biat.<br />
prasiniza) M. adnata; 245 M. prasina; 246 M. elachista; 248, 249 M. melaena; 269 {Bil. trisepta) M.<br />
peliocarpa; 270 {Bil. trisepta) M. nitschkeana; 307 M. misella; 357 {Bil. trisepta) M. peliocarpa; 407 M.<br />
melaena; 482 {Bil. trisepta) M. peliocarpa.<br />
BOHLER, Lich. Brit.: 85 (L. viridescens) M. lignaria.<br />
BRITZELMAYR, Lich. exs.: 174 {Biat. prasiniza var. laeta) M. adnata; 175 {Biat. glomerella) L.<br />
viridescens (M); 208 M. misella; 310 {Biat. syno<strong>the</strong>a) L. turgidula + Lee. sp. (H); 464 {Biat. syno<strong>the</strong>a) M.<br />
denigrata + C. nigroclavata (H); 599 {Biat. syno<strong>the</strong>a) Arthonia vinosa (H); 829 {Bil. milliaria f. nigrita)<br />
M. nitschkeana; 846 M. cinerea; 946 M. melaena; 961 (L. assimilata f. irrubata) L. hypnorum + L. limosa<br />
(H).<br />
CLAUDEL & HARMAND, Lich. Gall. : 43 M. lignaria; 89 M. nitschkeana; 445 M. misella.<br />
CROMBIE, Lich. Brit.: 174 (L. misella) M. misella +/or M. prasina.<br />
CUMMINGS, Decad. N Amer. Lich. Ed. I: 302 M. lignaria; 355 M. prasina. Ed. II: 232 M. lignaria.<br />
ELENKIN, Lich. Ross.: 189 M. misella.<br />
Erb. Critt. Ital., ser. I: 198 (C. syno<strong>the</strong>a) C. nigroclavata (UPS).<br />
FELLMAN, Lich. Arct.: 159 M. melaena; 164 (L. assimilata f. alpestris) M. incrassata, + a little M.<br />
crassipes in example in H; 165 (L. assimilata) M. crassipes; 166 (L. limosa) M. incrassata.<br />
FLAGEY, Lich. Franche-Comte: 137 {Bi. syno<strong>the</strong>a) Lee. sp. + Lecidella ? euphorea (UPS).<br />
Flora Hung, exs.: 714 M. melaenida.<br />
FLOTOW, Lich. exs.: 129A M. lignaria; 129B (L. milliaria b. ma/o/- - according to Lynge) Mycoblastus<br />
affinis (UPS); 129C (L. milliaria var. lignaria - according to Lynge) M. melaena; 129D (L. milliaria) C.<br />
globulosa (UPS); 129E M. lignaria; 130 M. turfosa; 131 M. lignaria; 171A M. sylvicola.
LICHEN GENUS MICAREA IN EUROPE 205<br />
FRIES, Lich. Suec: 29 M. lignaria, + some M. denigrata in example in UPS; 98 M. denigrata, + some M.<br />
lignaria in example in E; 212A, B M. melaena.<br />
HANSEN, Lich. Groenl.: 172 (L. assimilata) L. stenotera (BM); 247 (L. assimilata) L. limosa (BM).<br />
HARMAND, Guide Elem. Lich.: 91 M. lignaria.<br />
HARMAND, Lich. Loth.: 810 (Lecidea vernalis, and f. prasina) mixture from several localities, example<br />
in ANGUC has Bi. (? Moelleropsis) humida, C. sphaeroides, L. icmalea and L. vernalis, that in UPS has<br />
C. sphaeroides and L. vernalis; 838 M. denigrata; 852 M. lignaria; 853 M. nitschkeana.<br />
HAVAAS, Lich. Norv.: 139 M. subviolascens; 555 M. lignaria; 571 (L. sylvicola var.) M. lutulata; 694 M.<br />
subviolascens; 710 M. subviolascens.<br />
HAVAAS, Lich. Norv. Occid.: 269 M. subviolascens.<br />
HEPP, Flecht. Eur.: 14 M. denigrata; 20 {Bi. lignaria) M. nitschkeana; 21 (Bi. cinerea) M. cinerea, or M.<br />
nitschkeana; 278 M. prasina; 284 M. peliocarpa; 285 (Bi. lignaria var. milliaria) M. peliocarpa; 504 M.<br />
melaena; 510 M. peliocarpa; 524 {Bi. asserculorum) S. umbrinum (E).<br />
HEPP, Zurich: 206 {L. cinerea) M. peliocarpa; 210 (L. syno<strong>the</strong>a) Lecania cyrtella agg. (BERN); 224 M.<br />
prasina.<br />
HERTEL, Lecid. exs.: 34 M. lignaria; 54 M. ? bauschiana.<br />
JOHNSON, N. Engl. Lich. Herb.: 373 M. denigrata; 375 (L. turneri; listed under Bil. lignaria by Smith<br />
(1911, 1926)) Bacidia sabuletorum + Toninia lobulata (BM); 376 M. melaena; 434 L. sylvicola var.<br />
hellbomii) M. sylvicola, or M. bauschiana; 453 M. lignaria; 504 M. bauschiana.<br />
KALB, Lich. Neotropici: 22 M. lignaria (M. nigrata); 186 M. lignaria.<br />
KAVINA & HILITZER, Crypt. Cech.: 269 M. sylvicola.<br />
KORBER, Lich. Sel. Germ.: 12 M. turfosa; 75 M. sylvicola; 133A {Bil. lignaria) M. peliocarpa; 133B M.<br />
peliocarpa; 137 {Bi. denigrata) L. (Lecidella) alba (L); 250 M. prasina.<br />
Krypt. Exs. Vind.: 165 M. peliocarpa; 362 M. melaena; 658 M. lignaria; 1232 M. nitschkeana; 1532 M.<br />
misella; 2061 (C prasiniza var. prasinoleuca) Dimerella diluta (BM, BM ex K, M); 2268 M. assimilata;<br />
2561 (C denigrata) C. globulosa (BM, BM ex K); 3153 M. denigrata; 3154 M. melaenida; 3651 (L.<br />
misella) M. denigrata; 4214 M. misella; 4858 M. denigrata.<br />
KUTAK, Lich. Bohem. : 205 (L. asserculorum) M. denigrata; 310 M. prasina; All M. lignaria, + a little M.<br />
leprosula in example in O; 516 M. denigrata; 517 M. denigrata.<br />
LARBALESTIER, Lich. Caesar. Sarg.: 83 (L. arceutina var. hypnaea) M. lignaria; 84 M. sylvicola.<br />
LARBALESTIER, Lich. Herb.: 29 P. clavulifera; 68 M. bauschiana; 223 M. lutulata; 227 (L. polioides)<br />
M. tuberculata, but example in LIV is L. monticola;212 M. lignaria; 304 M. sylvicola; 305 M. sylvicola or<br />
M. bauschiana; 347 M. peliocarpa.<br />
LEIGHTON, Lich. Brit.: 120 (Bi. uliginosa) M. melaena; 210 M. lignaria; 238 (L. milliaria var. terrestris)<br />
M. lignaria, or M. peliocarpa, example in MANCH has a little M. leprosula; 386 M. lignaria; 388 (L.<br />
sabuletorum var. milliaria) M. botryoides.<br />
LOJKA, Lich. Hung.: 60 M. cinerea; 61 M. lignaria; 134 M. peliocarpa.<br />
LOJKA, Lichenoth. Univ.: 29-31 M. prasina; 137 M. nitschkeana; 233 M. lithinella.<br />
MAGNUSSON, Lich. Sel. Scand.: 134 M. prasina; 208 M. leprosula; 282 M. turfosa; 340 M. nitschkeana.<br />
MALBRANCHE, Lich. Norm.: 287 (L. sphaeroides var. lignaria) M. nitschkeana; 387 (L. sphaeroides<br />
var. melaena) M. denigrata.<br />
MALME, Lich. Suec: 20 M. rhabdogena; 21 M. elachista; 22 M. anterior; 23, 24 M. prasina, 25 M.<br />
nitschkeana; 26 M. eximia; 27 M. melaena; 28 M. contexta; 125 M. lithinella; 145 M. denigrata; 169 M.<br />
peliocarpa; 199 M. sylvicola; 216 M. assimilata; 288 M. lignaria; 362 M. crassipes; 365 (M. denigrata var.<br />
pyrenothizans) M. misella; 865 M. turfosa.<br />
MIGULA, Crypt. Germ., Aust., Helv.: 1 (5/7. milliaria) M. lignaria +lox M. leprosula, ± M. nitschkeana;<br />
132 M. denigrata; 226 M. lignaria.<br />
MOUGEOT & NESTLER, Stirpes Crypt. Vog.-Rehn.: 1329 (L. melaena) an example in E includes a little<br />
A/, melaena, o<strong>the</strong>r collections in BM (2 sets) , DBN and M (3 sets) contain a selection from <strong>the</strong> following,<br />
Lee. spp., L. granulosa agg., L. turgidula, L. sp., M. denigrata; 1430 M. lignaria.<br />
MUDD, Lich. Brit.: 156-8 M. lignaria; 159 M. melaena; 164 (L. prasina) L. viridescens (E, BM, M); 175<br />
M. sylvicola.<br />
NORRLIN & NYLANDER, Herb. Lich. Fenn.: 145 (L. sylvicola) L. conferanda (BM, H, M); 177 M.<br />
denigrata; 180 M. melaena; 182 (L. pelidna) M. intrusa [some examples may prove to be S. umbrinum];<br />
194A, B M. crassipes; 314 M. elachista, one example H is B. hegetschweileri; 319A, B (L. milliaria var.<br />
ternaria) M. lignaria; 321 M. turfosa; 724 M. elachista; lAA M. misella; 745 (L. asserculorum) M.<br />
denigrata; 763, 764 (L. tuberculata) M. sylvicola.<br />
OLIVIER, Lich. Orne: 237 M. melaena; 344 A/, lignaria.<br />
PISUT, Lich. Slov.: 156 M. //gnana.
206 BRIAN JOHN COPPINS<br />
POELT, Lich. Alp.: 22 (L. misella) M. denigrata.<br />
RABENHORST, Lich. Eur.: 224 {Bi. turfosa) L. uliginosa (BM ex K); 322 M. lignaria, example in M has<br />
some M. peliocarpa; 582 {Bil. lignaria) M. nitschkeana; 583 M. nitschkeana; 603 {Bil. syncomista) M.<br />
lignaria; 626 M. denigrata; 648 {Bi. bauschiana) M. bauschiana, or M. tuberculata; 675 (L. sylvicola) L.<br />
erratica (BM, M); 676 M. prasina; 733 M. prasina.<br />
RASANEN, Lich. Fenn. exs.: 489 {Bil. nitschkeana) B. naegelii (LD); 512 M. tuberculata; 642 M.<br />
nitschkeana; 651, 652, 653 M. prasina; 672 (L. sylvicola) M. sylvicola, +/or M. tuberculata, +/or M.<br />
lutulata; 822 (C elachista f. simplicata) L. turgidula (BM ex K); 963 M. melaena.<br />
RASANEN, Lichenoth. Fenn.: 137 (C. elachista f. simplicata) B. igniarii (BM ex K); 343 M. peliocarpa;<br />
344 M. melaena; 426 (C ///cw) M. globulosella.<br />
Reliq. Suza.: 42 M. misella.<br />
ROUMEGUERE, Lich. Gall.: 87 (L. vernalis var. milliaria) B. sabuletorum (BM, M); 193 (L. vernalis<br />
var. syno<strong>the</strong>a) M. lignaria; 231 M. melaena, but example in BM is mixture <strong>of</strong> a L. sp. and Lee. spp.; 232<br />
M. lignaria; 467 (L. melonida fmelaenidaj) Toninia aromatica (BM).<br />
SAMPAIO, Lich. Port.: 132 M. denigrata; 147 M. lignaria.<br />
SCHAERER, Lich. Helv.: 196 (L. sabuletorum var. lignaria) examples labelled 'Ad ligna decorticata in<br />
alpibus' are M. prasina, examples labelled 'In m. Belpberg' contain one, or a mixture, <strong>of</strong> <strong>the</strong> following,<br />
M. lignaria, M. melaena, M. peliocarpa and M. leprosula; 327 (Parmelia varia var. denigrata) L.<br />
(Lecidella) alba (BM ex K), or L. aff. cadubriae (E).<br />
SUZA, Lich. Bohem.: 131 M. sylvicola.<br />
THOMSON, Lich. Arct.: 12 (B. melaena) Toninia lobulata (LD).<br />
VAINIO, Lich. Bras.: 1420 M. misella; 1451 M. misella.<br />
VEZDA, Lich. Bohem.: 133 M. lignaria; 258 M. lignaria; 282 M. crassipes.<br />
VEZDA, Lich. Sel. exs.: 11 M. crassipes; 14 M. melaena; 90 M. prasina; 516 M. lignaria; 538 M. turfosa;<br />
706 (M. violacea) B. naegelii (BM); 858 (M. ternaria) M. lignaria; 957 (L. tuberculata) M. sylvicola; 1036<br />
M. lignaria; 1087 M. cinerea; 1088(M. ternaria)M. lignaria; 1134M. elachista; 1135 M. turfosa; 1341 (A/.<br />
tuberculata) M. sylvicola; 1342 M. peliocarpa; 1380 M. peliocarpa; 1430 M. denigrata; 1467 M. prasina;<br />
1595 (M. hemipoliella) M. prasina; 537 M. melaenida.<br />
WEBER, Lich. Exs.: 73 M. denigrata.<br />
ZAHLBRUCKNER, Lich. Rar. exs.: 110 M. nitschkeana; 152 (C. melanobola i.frullaniae =) Arthonia<br />
aff. exilis (BM); 174 (L. infidula) L. erratica; 175 M. prasina; 276 M. turfosa.<br />
ZWACKH, Lich. exs.: 121 M. lignaria; 122 (L. milliaria Fr. var. ?) M. elachista; 276 M. peliocarpa; 219<br />
(Bi. gelatinosa var. minorl) M. bauschiana; 279B M. bauschiana; 394 M. denigrata; 416 M. prasina; All<br />
(Bil. nitschkeana) S. umbrinum + S. chlorococcum (UPS); 470 M. nitschkeana [two labels: 'Bilimbia<br />
nitschkeana, bei Miinster in Westfalen'; or 'Lecidea nitschkeana, bei Dolbrueck in Kreise Paderborn<br />
. . .', sometimes labelled '470 bis\ example in M. also has M. denigrata]; 534 M. nitschkeana; 535 (L.<br />
latens) M. sylvicola; 587 M. nitschkeana; 590 M. lithinella; 591 A, B, 592A-E, 593A-C M. prasina; 594A,<br />
B, 595 M. bauschiana; 596 (L. latens) M. sylvicola; 597 M. sylvicola; 656 M. prasina; 657 M. melaena;<br />
675 M. melaena; 118 (L. cinerea) L. symmicta agg. (UPS); 780 (L. trachona) M. sylvicola; 897 M.<br />
peliocarpa; 898 M. cinerea; 900 M. turfosa; 919 M. sylvicola; 1085 M. misella.<br />
Examples <strong>of</strong> <strong>the</strong> following have not been examined by me; names taken from Lynge (op. d/.).<br />
BRITZELMAYR, Lich. exs.: 838 {Bil. lignaria); 890 {Bit. trisepta). FLOTOW, Lich. exs.: 112 (L. syno<strong>the</strong>a ?).<br />
HARMAND, Lich. Loth.: 834 (L. denigrata). KORBER, Lich. Sel. Germ.: 343 {Bit. syncomista). LARBALESTIER,<br />
Lich. Herb.: 139 (L. micrococca). OLIVIER, Lich. Orne: 264 (L. nitschkeana). ROUMEGUERE, Gen. Lich. exs.: 36<br />
(L. vernalis var. syno<strong>the</strong>a); 51 (L. vernalis var. milliaria). No examples <strong>of</strong> this exsiccata have been traced; it is almost<br />
certain that it was never distributed (P. M. J0rgensen, in litt.). SCHADE, STOLLE & RIEHMER, Lich. Sax.: 256 {B.<br />
trisepta). ZWACKH, Lich. exs.: 131 (L. glomerella and L. confluens).<br />
Acknowledgements<br />
My sincerest thanks go to Mr P. W. James and Dr F. Rose for <strong>the</strong>ir encouragement, guidance and many<br />
kindnesses, not only during <strong>the</strong> present study, but also from <strong>the</strong> time <strong>of</strong> my first interests in hchenology in<br />
1965. For fur<strong>the</strong>r help and advice in those earher years I here take <strong>the</strong> opportunity <strong>of</strong> extending my thanks<br />
to Dr D. L. Hawksworth, Mr J. R. Laundon, Dr M. R. D. Seaward, Dr T. D. V. Swinscow, and Mr A. E.<br />
Wade.<br />
I am much indebted to <strong>the</strong> curators and owners <strong>of</strong> <strong>the</strong> herbaria listed in 'Materials'; especially as <strong>the</strong>ir<br />
searches for relevant.specimens and processing <strong>of</strong> loans have <strong>of</strong>ten been long and arduous procedures.<br />
Fur<strong>the</strong>rmore, many <strong>of</strong> <strong>the</strong>se curators and collectors have provided me with much supplementary<br />
information on localities, habitats, and historical background. Special thanks are due to Miss F. J. Walker<br />
A
LICHEN GENUS MICAREA IN EUROPE 207<br />
<strong>of</strong> <strong>the</strong> <strong>British</strong> <strong>Museum</strong> (<strong>Natural</strong> <strong>History</strong>) for her efficient handUng <strong>of</strong> my many 'urgent' requests for<br />
herbarium and bibliographical information.<br />
Recognition for <strong>the</strong>ir helpful comments and suggestions on various nomenclatural and taxonomic<br />
problems pertaining to Micarea and related groups goes to Mr B. L. Burtt, Pr<strong>of</strong>. Dr H. Hertel, Dr H.<br />
Kilias, Pr<strong>of</strong>. P. M. J0rgensen, Pr<strong>of</strong>. Dr J. Poelt, Pr<strong>of</strong>. R. Santesson, Dr V. Wirth, and Dr A. Vezda.<br />
Much advice and assistance has come from many <strong>of</strong> my colleagues at <strong>the</strong> Royal Botanic Garden,<br />
Edinburgh; in particular, I would like to thank Pr<strong>of</strong>. D. M. Henderson, <strong>the</strong> Regius Keeper, and Mrs<br />
F. M. Bennell, Mr A. P. Bennell, Mrs N. M. Gregory, Dr R. Watling, Mrs J. M. Woods, Mrs H. Hoy, Mrs<br />
D. A. Morrison, Mr M. V. Ma<strong>the</strong>w, Mr D. G. Long, and <strong>the</strong> staff <strong>of</strong> <strong>the</strong> typing section.<br />
All <strong>the</strong> people and organizations to whom I am grateful for assistance with field excursions are too<br />
numerous to mention individually. However, special thanks go to Dr L. Tibell for arranging and leading<br />
me on a tour <strong>of</strong> mid- and north Sweden in 1977, and to <strong>the</strong> Nordic Lichen Society (especially Mr U.<br />
S0chting and Mr K. Ramkaer) for inviting me as guest visitor to <strong>the</strong>ir excursion in north Jylland in 1979.<br />
The following funds and institutions are thanked for financial support <strong>of</strong> field-work: Nature Conservancy<br />
Council, <strong>Natural</strong> Environment Research Council, Royal Botanic Garden, Edinburgh, and <strong>the</strong> World<br />
Wildlife Fund.<br />
Finally, I must thank my wife and family for tolerating <strong>the</strong> many sacrifices that <strong>the</strong>y have had to endure,<br />
especially during <strong>the</strong> preparation <strong>of</strong> <strong>the</strong> final manuscript.<br />
References<br />
Ahlner, S. 1948. Utbredningstyper bland nordiska barrtradslavar. Acta phytogeogr. suec. 22: i-ix, 1-257.<br />
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Index<br />
Accepted names are in roman and synonyms in italic. New names and principal references are in bold<br />
whilst an asterisk (*) denotes a figure or map. Some taxa mentioned incidentally (e.g. in lists <strong>of</strong> associated<br />
species) are not included.<br />
aberrans 185, 186<br />
abietina (Lecanactis) 67<br />
absistens (Bacidia) 88<br />
adnata 24, 29, 33*, 61, 64, 65, 66*, 67, 68*, 89,<br />
90, 93-95, 105, 108-110, 111*, 114, 204<br />
aene<strong>of</strong>usca (Lecidea) 198<br />
aeolotera (Huilia) 26<br />
aeruginosa 151<br />
aeruginosa (Lecidea) 198<br />
affinis (Mycoblastus) 204<br />
aggerata 186<br />
alabastrites 22, 23, 26*, 29, 32*, 34*, 64, 65, 85,<br />
89-91, 93-95, 99, 103, 110-113*, 122, 172<br />
alba (Lecidea, Lecidella) 205,206<br />
albella 121,122<br />
albicans 170<br />
albidolivens 170<br />
alnicola 151<br />
andesitica 128<br />
aniptiza 127<br />
anterior 24, 29, 32, 33*, 61, 64, 65, 69*, 90, 91,<br />
95, 100, 105, 108-110, 112-114, 136, 205<br />
antrophila 149<br />
aphanoides 138, 140<br />
apochroeella (Lecidea) 132<br />
arceutina (Bacidia) 197, 198<br />
arenarium (Microcalicium) 202<br />
arnoldiana (Bacidia) 67<br />
aromatica (Toninia) 206<br />
Arthonia 31,98<br />
Arthoniaceae 31<br />
Artho<strong>the</strong>lium 31<br />
asserculorum (Lecidea) 158, 159<br />
assimilata 22, 25, 29, 32, 35*, 61, 62, 64, 67,<br />
87-90, 95, 100, 104, 114-116*, 154, 186, 187,<br />
205<br />
atroviridis (Lecidea) 199<br />
Bacidia 88, 97<br />
bacidiella (Lecidea, Micarea) 135, 198<br />
bauschiana 24, 25, 29, 30*, 31, 36*, 61, 64, 71*,<br />
87, 89, 93, 95, 100, 105, 117-119*, 148, 149, 187,<br />
194, 204-206<br />
(A.<br />
beckhausii (Bacidia, Micarea) 20, 30, 88, 102,<br />
135, 189, 196, 198<br />
berengeriana (Lecidea) 115<br />
Biatora 97,112,197<br />
Biatorina 97<br />
biforme (Anisomeridium) 66<br />
biseptata \1A<br />
Bispora 29<br />
botryocarpa 192<br />
botryoides 24, 29, 32, 36* , 61 , 62, 64, 65* , 67,<br />
69*, 84, 88-90, 95, 99, 106, 108, 114, 118-121*,<br />
149, 156, 159, 163, 195, 205<br />
botryoides (Byssus) 174<br />
bouteillei (Catillaria) 197*, 199<br />
brasiliana 158<br />
Bryophagae 96, 196<br />
Bryostigma 31<br />
Buellia 31,61<br />
byssacea 174, 175<br />
Byssoloma 98, 202<br />
cadubriae (Lecidea) 206<br />
caesioatra (Lecidea) 30, 116<br />
calamophila 142<br />
callicarpa 117<br />
Caloplaca 31<br />
carbonicola 132<br />
carneoglauca (Bacidia) 67<br />
Catillaria 61,88,97<br />
catillarioides 151<br />
Chaeno<strong>the</strong>copsis 64<br />
chalybeia (Catillaria) 97, 199<br />
chlorococcum (Scoliciosporum) 91, 97, 206<br />
chrysophthalma (Chrysothrix, Micarea) 200<br />
Chrysothrix 98, 196<br />
cinerea 22-26, 29, 37*, 61, 63-66, 70*, 85, 87,<br />
88-91, 93-95, 99, 104, 108, 109, 112, 121-123*,<br />
124, 129, 141, 144, 166, 172, 204-206<br />
clavulifera (Psilolechia, Lecidea, Micarea) 105,<br />
107, 163, 193, 198, 201*, 202, 203*, 205<br />
Cliostomum 97<br />
coccinea (Micarea) 96, 200<br />
Collemaceae 20, 24
conferanda (Lecidea) 205<br />
confusula 183<br />
conglomerata 169<br />
contexta 24, 29, 31, 32, 38*, 61, 63, 64, 71*, 85,<br />
87, 88, 90, 91, 95, 100, 107, 124, 125, 134, 164,<br />
205<br />
contristans (Catillaria) 98, 115, 200<br />
contrusa 138<br />
crassipes 20, 28*, 29, 38*, 62-64, 67, 72*, 87-89,<br />
95, 96, 100, 106, 115, 116, 125, 126, 152, 187,<br />
204-206<br />
cupreola 170<br />
curvata 29, 38*, 64, 85, 87-89, 92, 95, 105, 126,<br />
127, 185<br />
cyanescens (Micarea) 200<br />
cyrtella (Lecania) 198, 205<br />
cyrtellina (Lecania) 66<br />
dedivitatum 174<br />
decrustata 158<br />
delicatula 121, 122<br />
demarginata (Lecidea) 149, 198<br />
denigrata 22, 23*, 24, 26, 29, 30, 32, 39*, 61, 64,<br />
66, 67, 73*, 84, 85, 87-90, 92-96, 99-102, 108,<br />
109, 127-130*, 131, 133-136, 148, 166, 182, 198,<br />
204-206<br />
diluta 112<br />
diluta (Dimerella) 205<br />
dilutiuscula 117<br />
discrepans 174<br />
discretula 111<br />
dufourei (Lecidea) 200<br />
ecrustacea Lamy 127<br />
ecrustacea Nyl. ex Vainio 181<br />
efflorescens (Lecidea) 85<br />
elachista 22, 23*, 29, 40*, 64, 84, 88, 90, 91, 95,<br />
99, 103, 105, 129, 131-133, 181, 204-206<br />
elegantior (Huilia) 26<br />
endocyanea 151<br />
endoleuca 64, 84-86, 93-95, 104, 122, 146, 147*<br />
epiphaeotera 151<br />
erratica (Lecidea) 67, 187, 198, 206<br />
erysibe (Lecania) 199<br />
erysiboides (Catillaria, Lecidea) 20, 114, 198<br />
euphorea (Lecidella) 204<br />
exigua 170<br />
exilis (Arthonia) 196, 197, 206<br />
eximia 24, 29, 40*, 61-64, 71*, 87, 90, 91, 95,<br />
100, 107, 124, 134, 164, 168, 205<br />
flotowii 186<br />
fraterculans 170<br />
friesiana 128<br />
friesiana (Argopsis) 85<br />
frullaniae (Catillaria) 196, 197, 206<br />
fucatus (Mycoblastus) 88<br />
fuliginea (Lecidea, Micarea) 96, 198<br />
fusca , spododes var . 165<br />
LICHEN GENUS MICAREA IN EUROPE 211<br />
fusca, syno<strong>the</strong>a f. 128<br />
fuscatula (Bacidia) 196<br />
Fuscidea 22,97,185<br />
fuscoatra (Lecidea) 97, 198<br />
fuscopurpurea (Arthonia) 31<br />
gelatinosa (Lecidea, Micarea) 198<br />
geomaea 142<br />
geophana (Steinia) 98<br />
glebosula VIA<br />
globifera 158<br />
globularis 49*, 158<br />
globulosa (Catillaria) 66, 204, 205<br />
globulosella 22, 23, 41*, 64, 74*, 85, 88, 90, 95,<br />
96, 99, 101, 134, 135, 166, 180, 189, 190*, 198,<br />
206<br />
glomerella 131<br />
gomphillaceum 63, 96, 143, 144<br />
graniforme (Cliostomum) 67<br />
granulosa (Lecidea) 87, 98, 198, 200, 205<br />
granvina 149<br />
Graphidaceae 31<br />
griffithii (Cliostomum) 67, 159, 199<br />
gymnomitrii (Bryomyces) 143<br />
hardangeriana 185<br />
hedlundii 24, 31, 40*, 64, 75*, 85, 88, 90, 91, 95,<br />
100, 102, 108, 114, 135-137, 140, 175<br />
hegetschweileri (Bacidia) 205<br />
hellbomii 186<br />
Helocarpon 96, 126<br />
hemipoliella 39*, 127, 131<br />
hemipolioides 170<br />
Herteliana 97<br />
Huilia 26, 97<br />
humida (Biatora, ?Moelleropsis) 205<br />
hylocomii (Micarea) 200<br />
hymenea (Pertusaria) 88<br />
hypnaea (Lecidea) 197, 198<br />
hypnorum (Lecidea) 31, 88, 115-116, 204<br />
hypocyanea 186<br />
hypoleuca 122<br />
hypopta (Lecidea) 132<br />
icmalea (Lecidea) 87 , 198 , 205<br />
igniarii (Bacidia) 206<br />
ilyophora 151<br />
incincta 186<br />
incrassata 22, 23, 25, 28*, 29, 42*, 61, 64, 67,<br />
72*, 88, 89, 95, 96, 100, 104, 115, 116*, 137, 138,<br />
204<br />
infidula 117<br />
infuscata 115, 137<br />
intrusa 20, 23, 25, 29, 32, 43*, 61, 64, 85, 87-89,<br />
95, 105, 138*, 139, 140, 184*, 185, 205<br />
irrubata 114<br />
juistense (Anisomeridium) 66, 67
212<br />
laeta 174<br />
lapiseda 170<br />
latens 192<br />
laxula 148<br />
Lecanora 61,88<br />
Lecanoraceae 98<br />
Lecidea 88, 97<br />
Lecideaceae 19, 31 , 71 , 87, 97, 98<br />
Lecidella 22,31,97<br />
leprosa (Vezdaea, Micarea) 200<br />
leprosula 22, 44*, 64, 85-87, 89, 90, 95, 99, 103,<br />
107, 140-142*, 172, 183, 205<br />
leucococca 170<br />
leucodontis (Bryostigma) 31, 196<br />
levicula (Lecidea) 87, 100<br />
lignaria 20, 22, 23*, 24, 27*, 29, 31, 45*, 61, 63,<br />
64, 66, 76*, 84-90, 92-96, 99, 104, 122, 126,<br />
129, 141, 142-145*, 146, 166, 170-172, 191, 192,<br />
197,201,204-206<br />
limosa (Lecidea) 98, 115-116, 199, 204, 205<br />
lithinella 24, 28*, 32, 46*, 61, 64, 85, 89, 92, 93,<br />
95, 100, 105, 118, 147, 148, 150*, 204-206<br />
littorella (Catillaria, Lecidea) 197* , 199<br />
livescens 170<br />
livida B. de Lesd. 165<br />
livida Korber 170<br />
lobariella (Dactylospora) 88<br />
lobulata (Toninia) 145 , 205 , 206<br />
longior 128<br />
lucida (Psilolechia) 202<br />
lutulata 24, 25, 29, 46*, 64, 65, 75*, 88, 89, 95,<br />
100, 106, 118, 120, 148-150*, 161, 187, 198,<br />
204-206<br />
lynceola 117<br />
major 128<br />
malmeana 134<br />
meizospora 143<br />
melaena 22, 29, 47*, 61-64, 77*, 85, 87-96, 100,<br />
103, 107, 124, 150-153*, 154, 165, 171, 187, 195,<br />
204-206<br />
melaenida 22, 24, 29, 48*, 64, 88-90, 92, 95, 100,<br />
106, 115, 154, 155, 195, 204-206<br />
melaeniza 24, 29, 47*, 64, 65, 69*, 85, 90, 91, 95,<br />
99, 106, 108, 114, 120, 155, 156, 159, 161, 164<br />
melanobola 24, 31, 47, 61, 64, 77*, 85, 88, 90, 95,<br />
100, 102, 156, 157, 175<br />
melanochroza 158<br />
Melanolecia 97<br />
melaphana 43*, 138<br />
meridionalis 154<br />
Metacapnodiaceae 67<br />
Micarea 20, 96-100<br />
Micareaceae 98<br />
microcarpa 165<br />
micrococca 173-176<br />
microspora 117<br />
milliaria 84, 142<br />
minima (Micarea) 200<br />
BRIAN JOHN COPPINS<br />
miniuscula (Bacidia, Micarea) 196, 198<br />
misella 24, 29, 31, 49*, 61, 64, 65, 67, 85, 88, 89,<br />
90, 94-96, 100, 102, 107, 114, 120, 156, 157,<br />
158-160*, 164, 175, 181, 182, 204-206<br />
monticola (Lecidea) 205<br />
moriformis 125<br />
muhrii 24, 29, 49*, 61, 64, 88-91, 95, 106, 148,<br />
149, 156, 160, 161<br />
myriocarpa 24, 29, 31, 32, 49*, 62, 64, 78*,<br />
88-90, 95, 105, 106, 148, 149, 161, 162*, 163<br />
myrtillicola (Bacidia) 196, 197*<br />
naegelii (Bacidia) 172, 206<br />
niger (Lichen) 200<br />
nigra (Micarea) 96, 200<br />
nigra, melanobola f. 174<br />
nigra, spododes var. 165<br />
nigella 24, 29, 31 , 50* , 61-64, 78* , 87, 90, 91 , 94,<br />
95, 100, 107, 108, 114, 120, 124, 134, 156, 159,<br />
163, 164*, 165<br />
nigrata Miill. Arg. (Bacidia, Micarea,<br />
Patellaria) 200,201,205<br />
nigrata Nyl. 143<br />
nigroclavata (Catillaria) 204<br />
nigrum (Collema, Placynthium) 200<br />
nitschkeana 22, 23, 30, 50* , 64, 66, 67, 85,<br />
87-96, 99-101, 129, 135-136, 165-167*, 171,<br />
172, 189, 204-206<br />
niveoatra (Opegrapha) 66, 67<br />
Nostoc 25,96,114,155<br />
obscurior 117<br />
occulta \1A<br />
ocelliformis (Lecidea) 199<br />
oculata (Pertusaria) 88<br />
olivacea 24, 29, 31, 50*, 62-64, 78*, 84, 89, 90,<br />
95, 100, 107, 124, 134, 164*, 165, 167, 168, 193<br />
osloensis 29, 31, 54*, 61, 64, 88, 89, 95, 106, 169,<br />
195<br />
pallida (Lecidea) 26<br />
panaeola (Huilia) 25, 140<br />
parissima 127<br />
paucula 149<br />
pauxilla (Lecidea) 199<br />
peliocarpa 22-25, 27*, 29, 51*, 61, 63-66, 79*,<br />
85-87, 89-91, 93-96, 99, 103, 104, 108, 109,<br />
111, 112, 122, 129, 141, 144, 149, 166, 169-173*,<br />
191,200,204-206<br />
perfurfurea (Pannularia) 198<br />
perpusilloides 196, 197*<br />
phaeobaea (Arthonia) 67<br />
Phyllopsora 22, 85<br />
Placynthiella 198<br />
poliococca 131<br />
poliococcoides 131<br />
poliodes 148<br />
polytrichi 53*, 80*, 96, 174, 176<br />
polytropoides 22 , 45 * , 143
populina (Lecidea, Micarea) 31,32<br />
praeviridans 127<br />
prasina Fr. 20, 23*, 24-26, 29, 30, 32, 52*, 53*,<br />
61, 64, 66, 80*, 85-96, 100, 102, 108, 109, 129,<br />
136, 148, 157, 173-176*, 177*, 178*, 179, 198,<br />
204-206<br />
prasina Tuck. & Mont. (Bacidia, Biatora) 203<br />
prasinata (Bacidia, Biatora) 203<br />
prasiniza 53*, 109, 173<br />
prasinoleuca 174<br />
pruinosum (Scoliciosporum) 30, 97, 180<br />
pseudoglomerella 128<br />
Psilolechia 98, 202<br />
pulverula 125<br />
pycnidiophora 23, 29, 54*, 64, 65, 67, 85, 89, 90,<br />
93, 95, 99, 103, 108, 114, 179, 180*, 181, 182<br />
pyrenothizans 111<br />
quercicola (Lecanora) 66<br />
radiata (Arthonia) 31<br />
recondita (Lecidea) 199<br />
relicta 154<br />
rhabdogena 24, 29, 31, 55*, 64, 85, 88, 90, 91, 95,<br />
99,105,133,164,181,182,205<br />
rhapidophylli (Bacidia) 196<br />
rhodosphaera (Catillaria) 197<br />
Rhopalospora 97<br />
rosei (Phyllopsora) 85, 146<br />
rosella (Bacidia) 97<br />
rusticella 117, 118<br />
sabuletorum (Bacidia) 145, 204-206<br />
salevensis (Bacidia) 171<br />
sanguineoatra (Lecidea) 115<br />
saprophila 151<br />
saxkola 142<br />
saxigena Hepp 169, 170<br />
saxigena Leighton 143<br />
scandinavica 192<br />
schadeanum (Scoliciosporum) 180<br />
schumannii 48*, 154<br />
Scoliciosporum 97, 98, 103, 140<br />
semialbula 128<br />
semipallens 117<br />
simplicata 131<br />
simplicior (Lecidea, Bacidia) 199, 200<br />
smaragdina 186<br />
sordidescens 173, 174<br />
sororians 131, 133<br />
spadicea (Arthonia) 67<br />
sphaeroides (Bacidia, Bilimbia, Catillaria,<br />
Lichen) 20, 112, 197, 198, 205<br />
spodiza 111<br />
spododes 165<br />
Sporacestra 203<br />
Steinia 98<br />
Stenhammarella 197<br />
stenotera (Lecidea) 98, 115-116, 205<br />
LICHEN GENUS MICAREA IN EUROPE 213<br />
Stereocauliscum 96<br />
Stigonema 25,96,114,115<br />
stipitata 23 , 29 , 55 * , 64 , 65 , 67 , 89 , 91 , 93-95 , 99<br />
103, 108, 114, 180*, 182<br />
stizenbergeri 151<br />
straminea (Lecanora) 192<br />
Strangospora 98<br />
subabietina (Lecanactis) 67<br />
subassimilata 125<br />
subdiscordans (Byssoloma) 98<br />
subfuscula (Bacidia) 192<br />
subinfidula 192<br />
subleprosula 20, 24, 31, 32, 56*, 64, 85-87, 89,<br />
94, 95, 99, 103, 107, 141, 182, 183, 191*<br />
sublivescens 111<br />
sublivida 186<br />
submisella 127<br />
subnigrata 22, 23, 29, 32, 57*, 61, 64, 67, 81*,<br />
88-90, 95, 105, 127, 139, 183, 184*, 185<br />
subrudecta (Bacidia) 201<br />
subviolascens 20, 22, 23, 25, 29, 57*, 64, 88, 89,<br />
95,100,101,115,185,186,205<br />
subviridescens 52*, 173, 174, 177<br />
subviridicans 201<br />
sylvicola 20, 23-25, 29, 31, 57*, 61, 62, 64, 66,<br />
82*, 87-90, 93, 95, 96, 100, 107, 118, 149, 152,<br />
186-188*, 193, 194, 199, 204-206<br />
symmicta (Lecanora) 30, 98, 206<br />
syno<strong>the</strong>a (Lecidea) 128, 199<br />
syno<strong>the</strong>oides 24, 32, 57, 58*, 64, 74*, 88, 90, 91,<br />
93-95, 100, 102, 157, 180, 188-190*<br />
tenebrosa (Schaereria) 88<br />
ternaria 22, 23, 29, 59*, 63, 64, 83*, 89, 95, 96,<br />
99, 104, 126, 144, 171, 190, 191*, 192<br />
terrestris (Lecidea) 199<br />
tetramera (Bilimbia) 112<br />
Toninia 22<br />
trachona (Bacidia) 67, 206<br />
Trapelia 199<br />
Trapeliaceae 199<br />
Trapeliopsis 199<br />
Tremolecia 97<br />
trisepta 170,171,200<br />
triseptataNyl. 143<br />
triseptata Hepp (Bacidia) 143<br />
triseptatula 170<br />
triseptatuloides 170<br />
tuberculata 24, 25, 29, 31, 59*, 61, 62*, 63, 64,<br />
82*, 84, 87, 89, 90, 95, 100, 107, 163, 168, 187,<br />
192, 193*, 194, 204-206<br />
turfosa Massal. 24, 29, 32, 60*, 62, 64, 88, 89, 95,<br />
96, 103, 106, 155, 194, 195*, 196, 199, 204-206<br />
turfosa Fr. 47*, 150<br />
turgidula (Lecidea) 30, 168, 204-206<br />
Tylothallia 97<br />
uliginosa (Lecidea) 169, 206<br />
umbrinum (Scoliciosporum, Bacidia,<br />
,
214<br />
umbrinutn - cont.<br />
Lecidea) 25, 92, 97, 127, 128, 138, 140, 205,<br />
206<br />
umbrosa 149<br />
vainioi 186<br />
vermicillifera (Opegrapha)<br />
vernalis (Lecidea, Biatora)<br />
verrucula 194<br />
vezdae (Bacidia) 196<br />
Vezdaea 24,30,98<br />
67<br />
20, 205<br />
BRIAN JOHN COPPINS<br />
vinosa (Arthonia) 204<br />
violacea 51*, 170<br />
violacea (Bacidia) 170<br />
viridescens (Lecidea, Micarea)<br />
viridis (Protococcus) 174<br />
vulgaris 128<br />
vulgata (Opegrapha) 67<br />
xylophila 143<br />
zsakii 22, 48*, 154<br />
\<br />
198,200,204,205