Bulletin of the British Museum (Natural History)

O' M' 

Bulletin of the 

British Museum (Natural History) 

A taxonomic study of the lichen genus 

Micarea in Europe 

Brian John Coppins 

Botany series Vol 11 No 2 29 September 1983

A taxonomic study of the lichen genus Micarea in 

Europe 

Brian John Coppins ](i, 

Royal Botanic Garden, Inverleith Row, Edinburgh, EH3 5LR 

Contents 

Synopsis 18 

Historical background 19 

Lecideaceae 19 

Micarea 20 

Materials 20 

Methods 21 

Field studies 21 

Light microscopy 21 

Thin-layer chromatography 21 

Nomenclature 21 

Morphology 22 

Thallus 22 

Phycobionts 25 

Cephalodia 25 

Apothecia: external features 26 

Apothecia: internal features 30 

Anamorphs (conidial states) 63 

Chemistry 84 

Lichen substances 84 

Pigments 87 

Ecology 88 

General habitats 88 

Specific habitats 90 

Distribution 93 

Britain 93 

Europe 94 

World 96 

The genus Micarea 96 

Intergeneric considerations 97 

Suprageneric considerations 98 

Infrageneric considerations 99 

Keys to species 100 

Guide to keys & identifications 100 

Key to European species 101 

Key to European species that may occur without apothecia 107 

The species 108 

1. M. adnata 108 

2. M. alabastrites HO 

3. M. anterior 112 

4. M. assimilata 114 

5 

. 

M. bauschiana 

6. M. botryoides 118 

7. M. cinerea 121 

8. M.contexta • 124 

9. M. crassipes 125 

10. M.curvata 126 

Bull. Br. Mas. nat. Hist. (Bot.) 11 (2): 17-214 

~ 

W. u-^^ 

H^ 

Issued 29 September 1983

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18 BRIAN JOHN COPPINS 

11. M.denigrata 127 

12. M. elachista 131 

13. M. eximia 134 

14. M. globulosella 134 

15. M.hedlundii 135 

16. M. incrassata 137 

17. M. intrusa 138 

18. M.leprosula 140 

19. M.lignaria 142 

20. M.lithinella 147 

21. M.lutulata 148 

22. M. melaena 150 

23. M. melaenida 154 

24. M. melaeniza 155 

25. M. melanobola 156 

26. M.misella 158 

27. M. muhrii 160 

28. M. myriocarpa 161 

29. M.nigella 163 

30. M. nitschkeana 165 

31 M. olivacea 167 

32. M. osloensis 169 

33. M. peliocarpa 169 

34. M. prasina 173 

35. M. pycnidiophora 179 

36. M. rhabdogena 181 

37. M.stipitata 182 

38. M. subleprosula 182 

39. M.subnigrata 183 

40. M. subviolascens 185 

41 M. sylvicola 186 

42. M. synotheoides 188 

43. M. ternaria 190 

44 M. tuberculata 192 

45. M.turfosa 194 

Excluded taxa 196 

Index to exsiccatae 203 

Acknowledgements 206 

References 207 

Index 210 

Synopsis 

A taxonomic revision is presented for the lichen genus Micarea in Europe, with special emphasis on those 

species occurring in the British Isles. In brief, this genus is here circumscribed to include most crustose 

Hchens with lecideine (biatorine) apothecia, a poorly defined excipulum (sometimes absent altogether) of 

radiating paraphysis-like hyphae, a non-granular epithecium, Lecanora-type, 8-spored asci, simple to 

multiseptate, hyaline ascospores, and a 'grass-green' phycobiont usually of the so-called 'micareoid' type. 

Comparisons and possible relationships with similar genera are discussed. Noteworthy discoveries made 

during this study include the finding of cephalodia in three species, dimorphic paraphyses in several 

species, and a wide array of anamorphic forms, with three species each found to have three conidial states. 

Type studies have been made for nearly all names known, as well as those suspected, to be referable to 

Micarea in its present, wider concept. Forty-five species are recognised, of which 31 are confirmed from 

Britain. Seven species are new to science: Micarea adnata, M. curvata, M. hedlundii, M. muhrii, M. nigella, 

M. olivacea, and M. myriocarpa Vezda & V. Wirth ex Coppins. Several described species are included 

in Micarea for the first time, and additional name changes are required for nomenclatural reasons; nine 

new combinations result: M. assimilata (Nyl.), M. crassipes (Th.Fr.), M. elachista (Korber) Coppins «fe 

R. Sant. , M. globulosella (Nyl) , M. intrusa (Th.Fr.) Coppins & Kilias, M. lignaria war. endoleuca (Leigh ton) 

M. melaenida (Nyl.), M. melanobola (Nyl.), and M. subviolascens (Magnusson). Several taxa are ex- 

,

LICHEN GENUS MICAREA IN EUROPE 19 

eluded from the genus and the new combinations, Psilolechia clavulifera (Nyl.) and Bacidia prasinata 

(Tuck.), are proposed. Keys for the identification of all the accepted European taxa are given. The taxonomic 

parts are preceded by an outline of the historical background to the study of Micarea, and details of 

materials and methods employed in this study. Detailed accounts of the morphology, chemistry, and 

ecology in the genus are provided , and a discussion of distributions is supported by maps for the British taxa. 

All Micarea species occur on acidic, nutrient poor substrata, and most are confined to cool-temperate, 

boreal, or oceanic regions; a few occur in arctic-alpine areas but the genus is poorly represented in 

dry, lowland, Mediterranean regions. Prior to this study, most of the species were little-known or 

misunderstood; clarification of their taxonomy has been achieved by paying particular attention to their 

anamorphic states, chemistry (including pigmentation), and detailed anatomy. Consideration of the 

distribution and ecology of the species has proved invaluable in ordering the taxonomic chaos which previously 

surrounded the notoriously variable species of the genus. 

Lecideaceae 

Historical background 

Until recently the circumscription of the Lecideaceae (and of the genera within it) had changed 

httle from that adopted by Zahlbruckner (1926). It included most Uchens with the following 

combination of characters: a crustose to squamulose thallus, a 'grass-green' phycobiont 

(excluding Trentepohlia and Phycopeltis) , ± immersed to sessile, disc-like apothecia without a 

thalhne margin, mainly colourless spores, and an absence of parietin (or related pigments) and 

(or) polarilocular spores. The principal genera in the family (e.g. Lecidea, Catillaria, Bacidia, 

Biatorella, Mycoblastus, Lopadium, Bombyliospora, and Toninia) were separated mainly on 

the basis of spore characters, i.e. size, septation, and number per ascus. This classification 

largely ignored many features which (according to modern mycological concepts) now merit 

careful consideration, although they were used to varying degrees for the delimitation of species 

or infrageneric categories above this rank. In brief, these features involve the structure of asci, 

excipular and hypothecial tissues, paraphyses and anamorphs, ontogeny, finer aspects of thallus 

structure, and nature and location of pigments and lichen substances. In addition, investigations 

of the phycobiont(s) and considerations of ecology and phytogeography often provide valuable 

supplementary information. However, the use of some of these features in an attempt to define 

more natural genera is not a purely recent phenomenon. Several lichenologists working in the 

1850s and 1860s made bold attempts in this direction. With regard to the Lecideaceae s. lat., two 

lichenologists deserving special mention are G. W. Korber (who introduced Lecidella, Lopadium, 

Pyrrhospora, Schaereria, Schadonia, and Steinia) and A. B. Massalongo (who introduced 

Catillaria, Psilolechia, Sarcosagium, Scoliciosporum, Strangospora, and Toninia). In the latter 

half of the 19th century lichenology came under the almost monarchical influence of William 

Nylander, whose simplistic generic concepts gained precedence over the more far-sighted works 

of Korber, Massalongo, and others. From the 1870s right up to the 1950s there were few 

attempts to reassess the generic concepts of Nylander or the slightly more complex, but no less 

artificial, system of Zahlbruckner. Between about 1929 and 1954 the French lichenologist M. G. 

B. Choisy resurrected many of the old and more or less forgotten genera, and created several 

new ones (e.g. Haplocarpon [= Huilia], Hypocenomyce, Trapelia, and Tremolecia). Unfortu- 

nately, Choisy's works made little impact at the time and it was not until the mid-1960s that 

lichenologists began to look more carefully at the delimitation of genera. Recent investigations 

have led to the reinstatement (although often with emendations) of many of these genera and 

many new genera have had to be described (e.g. Fuscidea, Herteliana, Melanolecia, Trapeliop- 

sis, Tylothallia, and Vezdaea). Most are included in the key to European lichen genera in Poelt 

& Vezda (1981). Despite the many advances made during the last 15 years, it will be several 

decades before a reasonably natural generic classification within the Lecideaceae s. lat. is 

achieved. The enormity of the task can be appreciated from the fact that Zahlbruckner 

(1921-40) accepted no less than 1450 species in the genus Lecidea alone! In addition to the high 

number of taxa involved , further problems arise from the locating and obtaining on loan suitable 

(including type) material, and the many difficulties in observing and interpreting many 

microscopical, morphological, and ontogenetic features.


Micarea 

The genus Micarea was first validly described in 1825 by Elias Fries in his Systerna orbis 

vegetabilis (see p. 96), and was placed in his 'Tribus Collemaceae' on account of its rather 

gelatinous thallus, although he noted that it had Lecidea-hke apothecia. The generic name was 

little used by most 19th century lichenologists, although it was accepted with the single species 

{M. prasina) by a few such as Korber (1855). Towards the end of that century, J. T. Hedlund 

submitted his doctoral thesis to the University of Uppsala. In this work (Hedlund, 1892) he 

adopted and emended the genus to include 20 species, and his circumscription of the genus is 

essentially the same as that accepted by Vezda & Wirth (1976) and myself, although many 

species have since been added. Hedlund's sagacious work was evidently too revolutionary for his 

time, and it did not achieve international recognition. It seems that Hedlund was disillusioned 

and had difficulties in finding a position. He turned to horticulture (especially dendrology) and 

then became an authority on Sorbus. Although Hedlund's obvious talents were not lost to 

botany, they were sadly lost to lichenology, especially the study of microlichens. 

Vezda & Wirth (1976) slightly expanded Hedlund's concept of Micarea by including Lecidea 

sylvicola and related species (all without 'micareoid' algae), and also Bacidia beckhausii. Apart 

from the exclusion of B. beckhausii, my own concepts are much the same, although I have 

emended the genus very slightly so as to include species such as Lecidea (Helocarpon) crassipes 

and Catillaria intrusa. 

The first species of Micarea to be described was Lecidea [Micarea] lignaria Ach. (1808). By 

1850 only seven of the currently accepted 45 European species were validly published. At about 

this time lichenologists began to make use of better quality microscopes and the additional 

characters they revealed. By the end of the nineteenth century 34 of the accepted species had 

been described. The European Micarea flora was increased by only one species (Lecidea 

subviolascens) between 1900 and 1960. In 1961 Vezda published Bacidia [Micarea] subleprosula, 

and as a result of the present studies (some in collaboration with Mr P. W. James, Dr A. 

Vezda, and Dr V. Wirth) a further nine species have been described. In the future a few 

additional species will probably be added to the European flora, but the greatest expansion 

within the genus will come from the study of extra-European collections; indeed, many 

European, undescribed, and described (in other genera) species are already known to me from 

parts of the world outside Europe. 

Materials 

During the course of this study I have examined about 3,000 specimens (c. 1800 from the British 

Isles) attributable to Micarea, plus a further c. 500 specimens which have been referred to other 

genera. Material has been received on loan from (or studied in) the following institutional 

herbaria: ABD, ANGUC, BEL, BERN, BG, BM, BON, C, DBN, DEE, DUKE, E, G, 

GLAM, GZU, H, HAMU, HBG, HEX, IMI, K (now in BM; cited as BM ex K), L, LD, LIV, 

LSR, M, MANCH, NMW, NWH, O, S, STD, STU, SUN, TUR, U, UPS, VER, WCR, WIS, 

WRSL; abbreviations according to Holmgren et al. (1981). In addition, numerous specimens 

have been received on loan from private herbaria. From the British Isles these are: Dr H. J. M. 

Bowen (Oxford), Dr R. W. M. Corner (Penrith), Mr I. P. Day (Carlisle), Dr U. K. Duncan 

(Arbroath; lichen herbarium recently gifted to E), Dr A. Fletcher (Leicester), Mr V. J. 

Giavarini (Parkstone, Dorset), Dr O. L. Gilbert (Sheffield), Mr R. Gomm (Taunton), Rev. G. 

G. Graham (Hunwick, Co. Durham), Mr A. Henderson (Leeds), Dr C. J. B. Hitch (Saxmundham, 

Suffolk), Dr P. D. Hulme (Aberdeen), Dr A. R. Pentecost (Royal Tunbridge Wells), Dr 

F. Rose (Liss, Hampshire; many specimens now in BM), Dr M. R. D. Seaward (Bradford), Mr 

J. F. Skinner (Southend-on-Sea), Dr P. B. Topham (Dundee), Mr R. G. Woods (Newbridge on 

Wye, Powys); and from elsewhere in Europe: Dr J. Hafellner, DrH. Mayrhofer, and Prof. Dr J. 

Poelt (all Graz, Austria), Mr L.-E. Muhr (Karlskoga, Sweden), Dr A. Vezda (Brno, Czechoslovakia), 

and Dr V. Wirth (Ludwigsburg, W. Germany). 

When studying a group of much misunderstood lichens it is a rewarding exercise to investigate 

folders of other superficially similar (but strictly unrelated) taxa, especially those that occur in


similar habitats. During this study numerous specimens of Micarea have been found in folders of 

widely misinterpreted ('dustbin') names, such as ' Bacidia sphaeroides' , 'Catillaria erysiboides' 

and Lecidea vemails' 

. 

Copies of the list of specimens examined have been lodged at BM, DBN, E, GZU, M, NMW, 

UPS and US. 

Field studies 

Methods 

I have attempted to observe as many as possible of the accepted species in their natural habitats 

because such experience is invaluable for the appreciation of environmentally controlled 

variation and ecological requirements. For this end I have made field studies in most parts of 

mainland Britain, and also S.E. Ireland, Denmark (N. Jylland), mid- and N. Sweden. In 

addition, I have collected Micarea specimens during earlier expeditions to France (Bretagne) 

and western Ireland. I have been successful in finding two-thirds of the European species, and all 

but two {M. asslmllata and M. subleprosula) of the 31 British species. 

Light microscopy 

Observations and measurements of external features were mostly made at x50 using a Vickers 

stereomicroscope equipped with a measuring eyepiece. Internal features were investigated with 

a Wild M20 microscope which was fitted with a drawing tube and a 2-5 x adapter for the drawing 

of spores and conidia etc. Most sections were made by hand with a razor blade, but some were 

cut by a freezing microtome. Sections were usually mounted in water, followed by (or directly 

in) 10% or 50% KOH(K), domestic bleach(C), 50% HNO3, or ammoniacal erythrosin (0-5g 

erythrosin in 100 ml 10% ammonia solution), but other mountants such as cotton-blue in 

lactophenol (LCB), congo red, and alcian blue were also used at times. Tests for amyloid 

reactions were made by mounting directly in Lugol's iodine solution (Ig iodine and 2g potassium 

iodide in 300 ml distilled water), or in this solution following treatment with 10% KOH. More 

permanent preparations were made by ringing mounts in LCB with nail varnish, or by mounting 

in polyvinyl alcohol (PVA) or cotton blue in PVA (see Omar etal, 1979). For further details on 

techniques see also 'Guide to keys and identifications' (p. 100). 

Thin-layer chromatography 

The t.l.c. techniques employed were those described by Walker & James (1980) which are based 

on the standard method of Culberson (1972) . For the purposes of routine analysis only two of the 

three basic solvent systems were found to be necessary (i.e. H.E.F. and T.D.A.). 

Nomenclature 

In the list of synonyms for a given species each entry begins with the oldest valid name 

(basionym) and is followed with later combinations included in Hedlund (1892), Smith (1911, 

1926), and James (1965^), and some others which have important nomenclatural implications. 

Zahlbruckner (1921-40) and Lamb (1963) should be consulted for additional combinations. The 

entries are listed chronologically, except for entirely invalid (or illegitimate) and misapplied 

names which are included at the end of the lists. 

Abbreviations of authors are according to Hawksworth (1980); those of journals to the third 

[1980] edition of Serial Publications In the British Museum (Natural History) Library; and those 

of books to Hawksworth (1974) or Stafleu & Cowan (1976-81). 

Apart from those newly described in recent years, few names in Micarea have been formally 

typified (according to Art. 7 of the Code); all cited lectotypes and neotypes are selected in this 

work, unless otherwise indicated. 

The nomenclature of lichens not treated in detail mainly follows Hawksworth et al. (1980) , but 

a few later changes are used. 

,


Morphology 

Thallus 

Thallus structure in Micarea is crustose and basically simple, but nevertheless, encompasses 

much interspecific and intraspecific variation. For the purposes of explanation thallus morphology 

can be roughly divided into five types. [Note: the discussion of the first type includes some 

general features applicable to all types.] 

(i) Areolate-type (Fig. lA-C) 

The term areolae (pi.) is used here to describe discrete, rounded (when viewed from above), 

flattened, or more usually convex (sometimes ± globose) portions of the thallus which have 

developed directly from the prothallus lying in or on the substratum. These should be 

distinguished from the (often angular) segments derived from the cracking of a previously 

continuous crust. The areolae in Micarea mostly range from the 0-06 to 0.2 mm diam, but are 

characteristically larger in certain species. Thalli comprising smaller, discrete, granular, soredia- 

Uke structures (goniocysts) are dealt with in the next category (ii). 

Species that have well-developed, convex areolae sometimes appear ± squamulose, e.g. M. 

assimilata, M. incrassata, M. lignaria (as in type of Lecidea trisepta var. polytropoides) , M. 

melaenida, and M. subviolascens. Catillaria zsakii (a synonym of M. melaenida) was placed in 

the squamulose genus Toninia Massal. by Lettau, but its thallus is not truly squamulose and it 

was excluded from Toninia by Baumgartner (1979). In M. subviolascens (and to a lesser extent 

in some other species) the areolae become confluent and the resultant crust becomes secondarily 

cracked into 'islands' (each containing several primary areolae), thus giving the thallus a rather 

squamulose appearance. The areolae in most species are whitish or pale to medium grey, and 

often tinged dull greenish, bluish or brownish. The areolae of M. lignaria var. endoleuca are 

characterically ivory or cream-yellow. More vivid yellow, citrine or orange thallus colourations 

are not known in Micarea. 

A few whitish, arachnoid, prothalline hyphae are sometimes visible in specimens with 

scattered areolae, but a thick prothallus as found, for example, in some species of Phyllopsora 

(Swinscow & Krog, 1981) is unknown in Micarea. The thallus in most Micarea spp. is effuse and 

wide-spreading, and even when thalli form small patches amongst other lichens, a delimiting 

hypothallus, such as found in many species of Fuscidea and Lecidella, is never formed. Small 

thalU forming ± circular patches amongst other lichens, suggesting parasitic behaviour, are 

characteristic of M. intrusa{p. 140). 

In vertical section the areolae usually lack a well-defined cortex, but their surface is sometimes 

covered by a thin (c. 2-10 (xm thick), hyaline amorphous layer which does not dissolve in 

concentrated KOH (Fig. lA, C). Such a layer is found in the areolae of, e.g. M. alabastrites, M. 

assimilata, M. cinerea, M. elachista, M. incrassata, M. lignaria, M. peliocarpa, M. subnigrata, 

M. subviolascens, and M. ternaria. Areolae with a similar external appearance, but without an 

amorphous covering layer, can be seen in well-developed specimens of, for example, M. 

denigrata, M. nitschkeana, M. globulosella, and sometimes A/, melaena (Fig. IB). The thallus of 

these species (and to a lesser extent that of M. elachista) is often invaded and disrupted by the 

dematiaceous hyphae of a fungal parasite(s), and non-lichenized algae, resulting in a dark, often 

blackish, scurfy crust. This phenomen on is best exemplified by M. melaena, which in Britain is 

rarely found with a healthy, well-developed thallus. By contrast, during my excursions in 

Sweden I found it to be rarely parasitised. Susceptibility to invasion by parasites or opportunists 

may be related to climatic factors, being increased in the British Isles by the overall weather and 

warmer conditions. It would appear, therefore, that the amorphous covering layer (never 

present in M. melaena) is an effective barrier against potential invaders, but whether or not this 

is the reason for its evolution is a matter for speculation. 

The nearest approaches to the development of a true cortex are found in M. elachista and M. 

melaenida. Sections of healthy areolae show an outer layer (c. 10-12 /xm and 12-20 /xm thick 

respectively) of compacted, evacuated hyaline hyphae (Fig. IC). The amorphous covering layer 

of M. elachista often partially disintegrates causing the areolae to exhibit a white-pruinose 

appearance, a feature not noted in any other species of the genus.


Fig. 1 Some thallus types in Micarea. A, areolate-type without cortex but with amorphous covering layer 

(e.g. LI lignaria). B, areolate-type without cortex or amorphous covering layer (e.g. M. denigrata). C, 

areolate-type with both cortex and amorphous covering layer (M. elachista). D, goniocysts (A/, prasina). 

Scale = 50 /Ltm. 

The outermost hyphae of areolae are frequently coloured or surrounded by a pigment , usually 

that which is found in the upper hymenium of the apothecia, or the pycnidial walls, of the given 

species; for example, the dilute olivaceous, K+ violet pigment in M. denigrata, M. elachista, M. 

globulosella, M. nitschkeana, and M. subviolascens, the green, K— , HNO3+ red pigment in M. 

cinerea, M. lignaria, M. peliocarpa, M. sylvicola, and M. ternaria, and the brown, K- , HNO3— 

pigment in M. subnigrata. The intensity of pigmentation is much dependent on exposure to light 

and pigment may be entirely absent in shade forms. Species that always lack pigment in their 

apothecia and pycnidia (e.g. M. alabastrites, M. pycnidiophora, amd M. stipitata) similarly lack 

pigment in the thallus. 

In the areolae of most species the algal layer is in direct contact with the substratum. However, 

a white medulla, devoid of algal cells, may be formed in the larger areolae of some species, e.g. 

M. incrassata, M. intrusa, M. lignaria, and M. subnigrata.


(ii) Goniocyst-type (Fig. ID) 

Several species have a finely granular thallus composed of discrete, ± globular structures, 

mostly c. 12-40 /am diam. These ecorticate granules consist of clustered algal cells intertwined 

and surrounded by short-celled hyphae, and are never protected by an amorphous covering 

layer. They are often seen to have short protruding hyphae, but they never have distinct spines 

as found in some species of Vezdaea (Poelt & Dobbeler, 1975). They are similar to soredia, but 

because they are the main component of the thallus and not derived from specialised structures 

(soralia) or eroding or disintegrating parts of a different thallus type (see (iii) below), the term 

goniocyst (Ozenda, 1963) seems the most applicable for them. 

In M. prasina, M. hedlundii, and M. melanobola the thallus is composed entirely of 

goniocysts, the first of which appear to arise directly from the prothalline hyphae; further 

development is presumably by a process of division or budding from existing goniocysts. M. 

botryoides has a 'primary' thallus of flattened granular-areolae and in a few specimens I have 

noticed goniocysts developing from these areolae, which then become obscured as the gonio- 

cysts proliferate. However, in most specimens of M. botryoides no areolae can be seen; either 

they have become obscured or, perhaps in some cases the thallus develops as goniocysts from the 

outset. 

In M. prasina and M. melanobola the outermost hyphae of superficial (exposed) goniocysts 

are often surrounded by the K+ violet pigment which also occurs in the upper hymenium of 

these species. When well developed, thalli composed of goniocysts have a ± gelatinous 

appearance when moist; this was the reason why Elias Fries originally placed the genus Micarea 

in his 'Tribus Collemaceae'. 

The thallus of M. synotheoides consists of small dark granular-areolae and is somewhat 

intermediate between the 'areolate-type'. The thallus of M. myriocarpa sometimes has a scurfy 

appearance but is often organized into small (10-15 /am diam) goniocyst-like granules. 

(iii) Sorediate-type 

This is represented by the apparently closely related M. leprosula and M. subleprosula. Their 

thalli are essentially of the 'areolate-type' but their areolae lack an amorphous covering layer 

and are very fragile (easily broken by touching with the point of a needle), often breaking 

down or eroding to form irregularly shaped, pale green or yellowish green soredial granules, 

c. 20-50 /Ltm diam. 

(iv) Smooth- or scurfy-type 

Included here are thalli which, although developed on the surface of the substratum, are not 

organized into discrete areolae or goniocysts. The thalh may be smooth, continuous to rimose, 

or irregularly scurfy-granular. Such thalli may be formed by species that normally have well 

defined areolae (e.g. M. lignaria and M. melaenida) or, by lignicolous species whose thalli are 

often endoxylic (e.g. M. anterior, M. nigella, and M. olivacea). The thalli of M. bauschiana and 

M. lutulata vary from smooth or rimose, to scurfy-granular, and lack areolae, whereas the 

similar M. lithinella, M. sylvicola, and M. tuberculata sometimes produce areolae. Another two 

species whose thalli fall within this category are M. adnata and M. turfosa. 

(v) Immersed-type 

The thallus of many lignicolous species is indistinct and developed below the surface of the 

substratum, which may have a bleached appearance (cf. M. muhrii). An endoxylic thallus is 

occasionally formed by species that more characteristically have superficial areolae, e.g. M. 

cinerea, M. denigrata, M. lignaria, M. muhrii, and M. peliocarpa. Conversely, M. melaeniza and 

M. misella are usually endoxyUc but forms with areolae are sometimes encountered. The thallus 

in M. anterior, M. nigella, and M. olivacea is often endoxylic, but a thin ± smooth crust is 

sometimes formed on the surface of the lignum. 

The thalli of M. contexta, M. eximia, and M. rhabdogena are invariably endoxylic. Sections of 

the substratum show small goniocyst-like clusters (c. 15-40 /am diam) surrounded and interconnected 

by pigmented hyphae. As in the relevant examples in thallus types (i-iii), and in M. 

turfosa in group (iv), the pigment involved is that found in the upper hymenium or pycnidial 

walls of the given species, viz. : dark green and K— in M. contexta and M. eximia, and olivaceous 

and K-f- violet in M. rhabdogena.


The species which at times grow on bryophytes usually have a superficial thallus, but even so, 

the thallus is often, in part, endocuticular. When on bark Micarea thalli are generally 

epiphloeodal, but at least partially endophloeodal thalli have been seen in M. cinerea and M. 

peliocarpa. 

Phycobionts 

Among the 45 species of Micarea accepted in this revision three types (? genera) of 'grass-green' 

phycobiont can be distinguished by LM observations on thallus squashes. Unfortunately the 

identities of these algae are unknown and they await critical studies by an algologist. In addition, 

two genera of blue-green alga may be involved in the formation of cephalodia (^.v.). 

The commonest alga in Micarea is that found in the type species {M. prasina) and was 

discussed at some length by Hedlund (1892, 1895). In the course of this study this alga is 

hereafter referred to as 'micareoid' (Fig. 2). Its cells are fairly regular in appearance, ± globose, 

thin-walled and c. 4-7 jxm diam. Reproduction within the thallus appears to be by a process of 

cell division in which the protoplast divides in two, followed by the laying down of a dividing wall 

which joins up with existing wall of the parent cell (Fig. 2A). No internal divisions into 

aplanospores as found in, for example, Myrmecia and Trebouxia, have been observed. Contact 

between the fungal hyphae and the algae cells is by intracellular haustoria (Peveling, 1974) 

which are readily seen at x 1000 (mounted in 10% KOH, followed by ammoniacal erythrosin). 

The haustoria may be peg-like (Fig. 2B, D-E), swollen to become clavate (Fig. 2G) or capitate 

(Fig. 2C, H), or spreading into a foot (Fig. 2F). In collapsed algal cells (Fig. 2H) the haustoria 

are intensively stained. A characteristic feature of the micareoid alga and its relationship with 

the mycobiont is that a hypha frequently becomes closely aligned along the dividing line of two 

separating algal daughter cells (Fig. 2D-E); the hypha concerned is often seen to penetrate one 

or both of the cells by haustoria. 

The second algal type is found in Micarea sylvicola and its presumed relatives M. bauschiana, 

M. lutulata, and M. tuberculata. The cells are thin-walled and irregular in size and shape, varying 

from 5-12 /xm diam when globose to up to c. 15 x 10 /xm when ± ellipsoid. I am undecided as to 

their mode of reproduction within the thallus. The large number of cells at the lower end of the 

size range is suggestive of aplanospore formation, but I have never observed a mother-cell 

undergoing division. Haustorial penetrations by the mycobiont hyphae have been detected, but 

they are never as distinct as those involving micareoid algae or the phycobiont of M. intrusa (see 

below). 

The third algal type is confined to M. intrusa, and has large globose cells, 7-21 /^m diam, with 

thick hyaline walls c. l-lfxm thick. Cell division within the thallus seems to be by a process of the 

division of a mother-cell into three or four daughter-cells (i.e. 'protococcoid' division). Many of 

the cells are clearly seen to be deeply penetrated by a haustorium (Fig. 55). This phycobiont 

looks very similar to that of Scoliciosporum umbrinum and may well be identical with it. 

Cephalodia 

For reviews of the morphological, taxonomic, and physiological aspects of cephalodia see Jahns 

(1974), James & Henssen (1976), and Millbank (1976). 

Cephalodia have been found in three species of Micarea: M. assimilata, M. incrassata and M. 

subviolascens . The first two species have thalli composed of verrucose areolae with a micareoid 

phycobiont. However, sections of their thalli reveal the presence of ± globose structures (c. 

200-600 /xm diam) containing a blue-green alga of the genus Nostoc. In many cases these 

structures are visible externally and closely resemble the areolae except that they are brown, and 

usually darker, in colour. Internally they consist of numerous ramifying fungal hyphae (presumably 

belonging to the Micarea) and dense masses of Nostoc cells which have lost their normal 

(when free-living) filamentous arrangements; and I am in no doubt that these structures are 

cephalodia. I am less certain of the status of the more loosely organized clusters of Stigonema 

which are sometimes associated with the same two Micarea species, and also M. subviolascens. 

However, the Stigonema filaments are, at least partially, disrupted and fungal hyphae are 

present.


A B 

Fig. 2 'Micareoid' phycobiont of Micarea alabastrites and its relationship with mycobiont hyphae ; see text 

for further details. Scale = 10 /xm. 

The three above-mentioned Micarea species are morphologically closely related and occur in 

exposed arctic-alpine situations. In such habitats the formation of cephalodia with a blue-green 

alga(e) which has the ability to fix atmospheric nitrogen, is of undoubted nutritional benefit to 

the mycobionts. Most morphological and physiological studies concerning cephalodia have 

involved macrolichens. Reports of cephalodia in crustose lichen are relatively few, although 

some are Hsted by James & Henssen (1976). Additional examples in the Lecideaceae include 

Huilia aeolotera, H. elegantior, H. panaeola (Hertel, 1977), and Lecidea pallida (Fries, 1874). 

Apothecia: external features 

As a rule the apothecia in Micarea are small to medium sized, convex to ± globose or 

tuberculate, and usually immarginate. However, this generalization encompasses considerable 

variation even within a single given species. With regard to shape, some species (e.g. M. 

denigrata and M. prasina) are very variable with apothecia that may be shallow-convex to 

convex-hemispherical (Figs 3A-B, 4A-B) and finally ± globose (Fig. 3D) or tuberculate 

(Fig. 3C). Apothecia of other species may be less variable: those of M. alabastrites and M. 

cinerea are broadly convex to hemispherical, sometimes tuberculate, but never ± globose; at the 

H


^>t. 

OQ 

LU 

3 3


3 00


Other extreme those of M. contexta, M. eximia, and M. myriocarpa are ± globose from the 

beginning and often tuberculate . The range of variabihty for most other species falls somewhere 

between these last two extremes. Apothecia are usually sessile (or partly hidden by adjoining 

areolae or goniocysts) but occasionally, as with M. lignaria, M. botryaides, and M. ternaria, they 

may be turbinate or short-stalked (due to vertical elongation of excipular and (or) hypothecial 

tissues), but only in M. crassipes and an undescribed species from New Zealand is this a constant 

feature (Figs 3E, 4B). Apothecia are usually immarginate, but in those species with a well 

differentiated excipulum (e.g. M. cinerea, M. denigrata, M. muhrii, M. peliocarpa, and M. 

ternaria), a faint marginal zone is sometimes apparent, especially in young apothecia. This zone 

is rarely raised above the level of the disc, but it is often paler in colour (especially when wet) and 

slightly more glossy. Even in M. crassipes, the only species whose apothecia regularly exhibit a 

distinctly raised marginal rim, this feature is soon occluded as the disc expands and becomes 

convex. 

The dimensions of apothecia in Micarea fall mainly within the range 0- 1-0-6 mm diam. In 

some species (e.g. M. denigrata, M. peliocarpa, and M. prasina) there is considerable variation 

in apothecial size, but in others it may be less so. Characteristically small apothecia (


Fig. 5 Sketch demonstrating a situation which M. bauschiana shows variation in the colour of its 

apothecia, from paUid (A), through pallid/brownish or pallid/blue-grey (B), to blue-black (C). Site: 

roadside bank in woodland in lower Glen Roy, Westerness (cf. Coppins 3492/3, E); the bank is c. 1 m 

high). 

M. denigrata, M. nitschkeana, and M. prasina. I know of no Micarea that has apothecia 

possessing pruina or, exhibiting a bluish (caesious) 'bloom' when wetted. Such features are due 

to the existence in the epithecium of minute colourless (or pale straw when dense) granules that 

dissolve in K, and are characteristic of several species often misidentified to Micarea, e.g. 

Bacidia beckhausii, Lecanora symmicta agg. , Lecidea caesioatra, L. turgidula, and Scoliciosporumpruinosum. 

Apothecia: internal features 

All tissues of the apothecia are bound by a weak gel-matrix which ± dissipiates in 10% KOH to 

clearly reveal the component parts (i.e. hyphae, paraphyses, and asci). In some species the 

apices of the paraphyses and/or the hyphae of the hypothecium are additionally cemented by 

dense pigment deposited in the matrix and/or on the hyphal walls, thus tending to impair their 

separation in K. The presence of the gel-matrix serves to separate species of Micarea from some 

superficially similar species of Vezdaea (Poelt & Dobbeler, 1975). 

Hymenium and epithecium 

The term 'epithecium' is here employed in its broad sense to refer to the upper part (usually c. 

3-15 /xm) of the hymenium where this differs in appearance from the remaining lower part(s).


This difference can be due to the much branched and entangled apices of the paraphyses, or 

(and) deposits of pigment in the gel-matrix, or on (or in) the walls of the paraphyses. The upper 

layer of the hymenium differentiated in such ways is perhaps more correctly termed the 

'epihymenium' (Poelt, 1974a) or 'pseudoepithecium' (Korf, 1973). In its strict (original) sense 

an 'epithecium' refers to a layer of branches of paraphyses that overtop the asci. Such a layer is 

approached in some species of Micarea and is best exemplified by M. contexta and M. 

melanobola. In Micarea the difference between an 'epihymenium' and an 'epithecium' is not 

clear-cut, and so I have chosen to employ the latter term which has been long and widely 

employed in lichenology. 

The height of the hymenium in Micarea species is sometimes difficult to measure accurately 

because the hymenium often merges rather imperceptibly into the hypothecium. Measurements 

are most accurately made by mounting thin sections in Lugol's iodine, in which the strongly 

amyloid (dark blue) hymenium contrasts strongly with the non-amyloid (or ± so) hypothecium. 

Even when accurately determined the height of the hymenium is rarely a useful character in 

distinguishing similar species. The overall range in height is about 23-90 /am, and in most cases is 

within 35-50 /xm. Shallow hymenia (rarely exceeding 35 /am) are characteristic of many of the 

species with small spores, viz.: M. hedlundii, M. melanobola, M. misella, M. myriocarpa, M. 

nigella, M. olivacea, M. osloensis, M. rhabdogena, and M. tuberculata. Likewise, tall hymenia 

are characteristic of species with large spores, and the tallest hymenium (65-90 /u,m) belongs to 

the species with the largest spores, namely M. subleprosula. The height of the hymenium 

sometimes increases with the age of the apothecium, and this is particularly true in M. 

bauschiana and M. sylvicola: small, yet mature, apothecia usually have a hymenium c. 40 /zm 

tall, but as the apothecium enlarges and increases in convexity, the hymenium often increases in 

height up to about 60 /xm; at the same time the paraphyses appear to 'stretch' and become 

thinner (especially in the lower part of the hymenium). 

The hymenium in Micarea never contains hyaline, crystalline inclusions that adhere to 

paraphyses (as found in some species of Lecidella, and many Graphidaceae) or numerous oil 

droplets (as found in several species of Buellia and Caloplaca). Epithecial granules that dissolve 

in K and often give the apothecia a pruinose appearance are also never found in Micarea. Minute 

granules of blue-violet (K+aeruginose) pigment, like those found in Lecidea hypnorum, are 

occasionally observed in the hymenium of M. contexta and M. lignaria. 

The colour of the hymenium and epithecium is an important character in the identification of 

Micarea species. A discussion of the various pigments involved is given in the sections on 

'chemistry'. The hymenium may be dilutely and ± evenly coloured throughout, or the 

colouration may be more intense in the upper part. In many species the hymenium is often 

intersected by dark vertical streaks, a feature usually due to dense pigment surrounding 

individual, or small fascicles of stout paraphyses (see p. 61). 

Asci 

The asci of Micarea species are clavate or cylindrical-clavate in shape and belong to the 

Lecanora-type of Honegger (1978). When mounted in Lugol's iodine (following treatment in 

10% KOH) the ascus is seen to have a non-amyloid wall surrounded by an amyloid outer-layer 

('fuzzy coat') and an internal staining amyloid apical dome (Fig. 6). Micarea has sometimes 

been placed in the Arthoniaceae (e.g. Lamb, 1953) but the ascus of members of that family does 

not have an amyloid outer-layer and the apical dome is not deeply amyloid. In a few species 

of Arthonia and Arthothelium I have observed a faint bluing in the part of the apical dome 

immediately adjacent to the ocular chamber. In some Arthonia species a tiny amyloid ring is 

apparent above the apex of the ocular chamber, and, despite statements to the contrary by 

Eriksson (1981), such a ring is found in the asci of ^. radiata (type species oi Arthonia Ach.) and 

A. fuscopurpurea. The same type of ring was also described for Bryostigma leucodontis by Poelt 

& Dobbeler (1979). Such aspects of ascus structure in the Arthoniaceae clearly merit more 

detailed investigation. The ascus structure in Micarea supports my opinion that the genus should 

be placed in the Lecideaceaes. str., at least for the time being. 

The asci of all European Micarea species are 8-spored. The North American Lecidea populina


Fig. 6 Ascus apex of Micarea alabastrites in optical section (LM); mounted in Lugol's iodine following 

pre-treatment in 10% KOH. A, young ascus. B, mature ascus. ac, apical cushion; aw, ascus wall; d, 

apical dome (tholus); oc, ocular chamber; ol, outer layer of ascus wall. Shading indicates intensity of 

amyloid reaction; note that in reality the apical cushion and ocular chamber are probably completely 

non-amyloid. Scale = 10 /xm. 

Miill. Arg. has 16-spored asci and was transferred to Micarea by Anderson & Carmer (1974). 

Unfortunately the type material of L. populina has been out on loan from H and not available to 

me during the course of this study. However, the species apparently occurs in Xanthorion 

communities and, if this is so, then it is unUkely to be a Micarea. 

Spores 

The wide variety of spore types found in Micarea can be seen in the selection of spores from all 

the European species illustrated in Figs 7-33. A few species (e.g. M. assimilata, M. contexta, and 

M. lithinella) have spores that are ± consistent in size, shape, and septation, but for many other 

species (e.g. M. anterior, M. botryoides, M. denigrata, M. prasina, and M. turfosa) these 

characters can be very variable even within the same ascus. The smallest spores are found in M. 

myriocarpa (5-5-8-5xl-5-2-5 jxm) and the largest ones are found in M. subleprosula (40-60X 

5-6-6 /Ltm). Spores may be simple or up to 7-septate, the variations within these hmits depending 

on the species. Spores with more than 7-septa are very rare, although a few 9-septate spores have 

been observed in M. subleprosula, and a collection (Coppins, 1834) of M. synotheoides has 

spores with up to 11 septa. Among the great range of spore shapes encountered, some may be 

broadly ellipsoid {M. subnigrata and M. intrusa) , 

but regularly globose spores do not occur in 

Micarea. 

Healthy spores are always thin-walled and colourless. However, the walls of old spores 

trapped within the hymenium sometimes become slightly thickened and pale straw coloured, or 

they may become impregnated with hymenial pigment. Spore walls always appear smooth (LM 

at X 1000) and are never surrounded by a gelatinous epispore. The cytoplasm of spores, asci, and 

ascogenous hyphae in M. intrusa is sometimes a dilute orange, turning purple-red in K.


Fig. 7 A, M. adnata (E - holotype). B, M. anterior (H-NYL 21655 - lectotype). Scale = 10 /xm. 

B


Fig. 8 M. alabastrites. A, (H-NYL 18656 - holotype). B, {Coppins 2588, E). Scale = 10 /ixm. 

B

h 

LICHEN GENUS MJCAREA IN EUROPE 35 

Fig. 9 M. assimilata. A, (H-NYL 16556 -lectotype). B, (Norway, Oppland, Fogstuen, 1857, Lindsay, E). 

Scale = 10 /xm. 

B


Fig. 10 A, M. bauschiana (M - lectotype). B, M. botryoides {Coppins 8429, E). Scale = 10 /xm.


I- H 

Fig. 11 M. cinerea {Coppins 2533, E). Scale = 10 /Ltm.


Fig. 12 A,M. contexta (S-\ectotype). B,M. crassipes (VezdaLich. Sel. 11,BM).C,A/. cMrvara (WRSL 

holotype). Scale = 10 fxm. 

H 

B 

-


f]r\ 

VJ u 

Fig. 13 M. denigrata. A, (UPS - lectotype). B, (H - lectotype of Lecidea hemipoliella). Scale = 10 ^tm. 

B


r\ 

U 

Fig. 14 A, M. elachista (L - lectotype). B, M. eximia (S - lectotype). C, M. hedlundii (UPS - holotype). 

Scale = 10 /xm.


Fig. 15 M. globulosella. A, (S - lectotype). B, (Czechoslovakia, Slovakia, Vysoke Tatry, 1879, Lojka, 

BM). Scaler 10 /am. 

y


Fig. 16 M. incrassata. A, (S - holotype). B, (Kerguelen, 1875, Eaton, E). Scale = 10 /u,m. 

B


Fig. 17 M. intrusa. A, (UPS - holotype). B, (BM - isotype of Lecidea melaphana). C, (Norway, 

Hordaland, Fjell, 1980, Skjolddal, BG). Scale = 10 /xm.


Fig. 18 M. leprosula (UPS-lectotype). Scale — 10)u,m.


Fig. 19 M. lignaria. A, (H-ACH 265 - lectotype). B, (TUR-VAINIO 21274- lectotype of Lecidea trisepta 

yar.polytropoides). Scale = 10 ixm. 

H


Fig. 20 A-B, M. lithinella; A, (M - lectotype) ; B, (Germany, Heidelberg, Zwdckh, H-NYL 19191). C, M. 

lutulata (Wales, Pembroke, Tycanol, 1980, James, BM). Scale = 10 ^tm. 

B


B 

Fig. 21 A, M. melaena (UPS - lectotype of Lecidea milliaria var. turfosa). B, M. melaeniza (S - holotype) 

C, M. melanobola (H-NYL 21614 - lectotype). Scale = 10 /Ltm. 

.

48 

BRIAN JOHN COPPINS 

Fig. 22 M. melaenida. A, (H-NYL 18843 - holotype). B, (Fl. Hung. exs. 714, BM - topotype of Catillaria 

zsakii). C, {^'RSL-\ec\o\.yptoi Catillaria schumanii). Scale = lOjLtm. 

B


Fig. 23 A, M. misella (H-ACH 52-holotype of Lecidea resinae subsp. globularis). B, M. muhrii (E 

holotype). C, M. myriocarpa (hb Wirth 6085 - holotype). Scale = 10 /u,m. 

B


Fig. 24 A, M. nigella (E - holotype). B, M. nitschkeana (Coppins 2426, E). C, M. olivacea (BM 

holotype). Scale = 10/u,m. 

B


Fig. 25 M. peliocarpa. A, (H-NYL 18716 - isotype of Lecidea violacea). B, (England, New Forest, Great 

Wood, Bramble Hill Walk, 1970, Coppins et al. , E). Scale = 10 />tm. 

B


Fig. 26 M. prasina. A, (UPS - lectotype). B, (H-NYL 19056 - lectotype of Lecidea subviridescens). Scale 

= 10 /um. 

B


Fig. 27 M. prasina. A, (H-NYL 21604 - lectotype of Lecidea prasiniza) . B, (GZU - holotype of Micarea 

polytrichi). Scale = 10 /xm. 

B


VJ 

Fig. 28 A, M. pycnidiophora (E - holotype). B, M. osloensis (UPS - holotype). Scale = 10 ju-m. 

B


r\ 

Fig, 29 A,M. rhabdogena (BM - isolectotype). B, M. stipitata (E - holotype). Scale = 10 /u-m. 

\j


Fig. 30 M. subleprosula (Mu/ir 4380, E). Scale = 10 /Am.


Fig. 31 A, M. subnigrata (H-NYL 19136 -lectotype). B, M. subviolascens (Havaas Lich. Exs. Norv. 139, 

BG). C, M. sylvicola (UPS - lectotype). D-E, M. synotheoides; D, (H-NYL 19101 - lectotype); E, 

(Coppins 3257, E). Scale = 10 ixm. 

Fig . 3 1 D-E on foliowing page 

B

58 


Fig. 31 - cont.


^ 

Fig. 32 A, M. tuberculata (O - lectotype). B, M. ternaria {Thomson 9188, DUKE). Scale = 10 /xm. 

r\ 

u


Fig. 33 M. turfosa. A, (VER- holotype). B, (Vezda Lich. Sel. 1135, BM). Scale = 10 /xm.


Paraphyses 

The paraphyses in Micarea are characteristically thin and branched. When measured (in 10% 

KOH, or ammoniacal erythrosin) at the mid-hymenium they may be very thin and only about 

0-6-1 fxm wide (e.g. M. anterior, M. botryoides, M. contexta, M. eximia, M. lithinella, M. 

misella, and M. prasina), or thin and c. 1-1-5 /xm wide (e.g. M. adnata, M. cinerea, M. denigrata, 

M. intrusa, M. muhrii, and M. peliocarpa), or relatively stout and about 1-5-1-8 fjun wide (e.g. 

M. assimilata, M. incrassata, M. lignaria, M. osloensis, and M. subnigrata) ; these measurements 

relate to paraphyses not coated in pigment. In old, much expanded, apothecia the paraphyses 

sometimes appear 'stretched' and thinner than normal (especially in the lower half of the 

hymenium) ; this phenomenon has frequently been observed in collections of M. bauschiana, M. 

sylvicola, and M. lignaria. In many species the paraphyses gradually widen towards their apices, 

but the apices are never regularly clavate or capitate. This widening is often enhanced by the 

deposition of closely adhering pigment which sometimes gives the appearance of a 'hood' (e.g. 

M. melaena) ; such coatings or hoods can be detached by gently boiling and then tapping sections 

or squash preparations in 50% KOH. In a few cases (e.g. M. melanobola) the pigment cannot be 

separated in this way and appears to be located within the walls of the paraphyses. Paraphyses 

with dark pigmented apical 'caps', like those found in Catillaria s. str. (Killias, 1980: 253), 

Buellia, and many species of Lecanora, are not known in Micarea. 

In all species of Micarea a large proportion of the paraphyses are branched, even if the 

branching is mainly confined to the epithecium. Species with sparingly branched paraphyses 

include M. assimilata, M. incrassata and M. lignaria. Anastomozing paraphyses have been 

observed in all the species, but often the anastomoses are ± confined to the lower third of the 

hymenium. The degree of branching and anastomosing is difficult to quantify for the practical 

purposes of identification, but this character can be useful when comparing collections micro- 

scopically. 

A similarly difficult character is the relative abundance of the paraphyses. The two extremes 

can be referred to as: (a) 'numerous' - large in number and immediately discernible when 

observing a mount in 10% KOH at x400; (b) 'scanty' - few in number and not immediately 

obvious when observed in the same way. In hymenia with scanty paraphyses the 'extra space' is 

taken up by hymenial gel or a higher proportion of asci, or a combination of both. The situation 

in most species lies somewhere between the two extremes. Furthermore, the relative propor- 

tions of hymenial gel and asci to paraphyses sometimes increases as the apothecium expands (cf 

the above example of M. bauschiana, M. sylvicola, and M. lignaria). An accurate assessment of 

the abundance of paraphyses is not usually essential for the routine identification of Micarea 

species, although it can be helpful when comparing difficult, convergent forms of M. denigrata 

and M. misella (see couplet 11 of the main key). 

In addition to the 'normal' paraphyses described above, the hymenium of several species (e.g. 

M. bauschiana, M. botryoides, M. eximia, M. nigella, M. sylvicola, andM. tuberculata) contain 

scattered individuals, or small fascicles, of rather stout paraphyses. These 'paraphyses' are 

about 2-3 /xm wide and usually more distinctly septate than normal paraphyses. Furthermore 

(especially in species with a dark hypothecium), they are often coated in pigment throughout 

their length (Fig. 34), with the result that the hymenium is seen to be intersected by dark vertical 

streaks. They are mainly found in species with 'scanty' paraphyses, and appear to extend deep 

into the hypothecium. The elucidation of their true status and function awaits detailed 

ontogenetic studies, but it is possible that they have a strengthening, spacing or protective 

function during the maturation of the hymenium from the primary corpus. 

Hypothecium 

The area of tissue lying below the hymenium, or between the hymenium and the excipulum (if 

present), is generally referred to by lichenologists as the 'hypothecium'. In many groups of 

lichenized discomycetes (including the Lecideaceae) this area can be divided into an upper, 

usually narrow, layer containing mainly ascogenous hyphae, and a lower, often much deeper 

layer of structural tissue (hyphae gelatinised to various extents, according to genus or species). 

These two layers are often of different colour or colour intensity. Where the two layers are 

.


Fig. 34 Two types of paraphyses in Micarea tuberculata. A, normal, non-pigmented paraphyses. B, stout, 

pigmented paraphyses. Scale = 10 /^m. 

distinct they have been referred to as the 'subhymenium' and 'hypothecium' respectively (e.g. 

Hertel, 19776). The term 'hypothecium' in this context has been substituted by many students of 

non-lichenised discomycetes by the term 'medullary excipulum' (Korf, 1973), and was called the 

'ental excipulum' by Eckblad (1968). In Micarea the two layers are well-differentiated in only a 

few species (e.g. M. crassipes); and in the present work 'hypothecium' refers to all ascocarp 

tissue lying below the hymenium, apart from the excipulum (where present). This same 

terminology was adopted in the recent study of Catillaria by Kilias (1981). 

The hypothecium in Micarea is composed of deeply staining (e.g. in LCB and ammoniacal 

erythrosin), swollen-celled ascogenous hyphae (c. 2-5 /am wide) and moderately staining, 

slender 'structural' hyphae (c. 0-7-2 fxm wide; slightly varying in width according to species), 

embedded in a gelatinous matrix. In species with a colourless or pale hypothecium the matrix is 

± dispersed in K and the hyphae are then clearly visible ( x 400) . In some species with a coloured 

hypothecium the pigment may be ± evenly distributed through the matrix and the hypthae are 

similarly distinct in K (e.g. M. eximia, M. olivacea, and M. turfosa). However, in the majority of 

species with a darkly coloured hypothecium (e.g. M. assimilata, M. botryoides, M. melaena, M. 

myriocarpa, M. nigella, M. sylvicola, and M. tuberculata) the pigment, although present in the 

matrix, exists as a strongly adhering coat around the hyphae, such that the hyphae appear 

thick-walled and broad (c. 1-5^ /xm). The pigment coatings tend to bind the hyphae together so 

that they are often indistinct in K. For the routine identification of Micarea species it is not 

essential to know the location of pigment (i . e . ± 

B 

H 

evenly distributed through the gel-matrix versus


tightly bound to hyphae) but such knowledge is sometimes of supplementary value, e.g. when 

comparing M. eximia versus M. nigella and M. olivacea versus M. tuberculata. 

A more important diagnostic character in Micarea is the colour of the hypothecium in water 

mounts, and the corresponding colour changes obtained by the addition of KOH and HNO3. A 

discussion of the pigments involved is given under 'chemistry'. 

The height of the hypothecium (in vertical section) is largely dependent on the overall size and 

(especially in species with a poorly developed excipulum) convexity of the apothecium. The 

measurements given in the species descriptions relate to normally developed, non-tuberculate 

apothecia. This character is often very variable for a given species and consequently of Uttle 

diagnostic value when comparing closely similar species. One exception to this is the case of M. 

contexta (20-90 /am) versus M. melaena (80-160 /xm). 

In Lugol's iodine the hypothecial tissues are non-amyloid, although there is sometimes a faint 

bluing in the vicinity of ascogeneous hyphae (especially in the upper part of the hypothecium). 

Excipulum 

The size and distinctiveness of the excipulum ('ectal excipulum') in Micarea varies greatly 

according to species and the age of the apothecium. In species such as M. cinerea, M. crassipes 

(Fig. 4B), A/, peliocarpa, and M. ternaria the excipulum is sufficiently well developed that their 

young apothecia are often weakly or distinctly (M. crassipes) marginate in outward appearance. 

However, even when well developed and initially distinct, the excipulum may become reflexed 

and ± occluded as the apothecium expands and increases in convexity (Fig. 3A-B) or becomes 

tuberculate (Fig. 3C). In many species the excipulum is always extremely reduced or absent 

(Fig.4A). 

When present, the excipulum is composed of outwardly radiating branched and anastomosing 

hyphae that ± separate in K. The hyphae closely resemble paraphyses, but are usually more 

dense and more richly branched. With markedly convex apothecia it can be difficult to 

distinguish between reflexed portions of the hymenium and what might be an excipulum. In such 

cases the excipulum (if present) can be identified in good thin sections by the absence of asci and 

a negative (non-amyloid) reaction to Lugol's iodine. The excipulum often differs in colour or 

colour intensity from the hymenium, although a similar colour difference may sometimes be 

shown by reflexed parts of the hymenium. 

In M. crassipes and rare forms of M. lignaria ('f. gomphillaced') the excipular and hypothecial 

tissues become vertically extended to form a stipe (Figs 3E, 4B). 

Anamorphs (conidial states) 

With a few noteworthy exceptions, such as Lindsay (1859, 1872) and Gliick (1899), the conidial 

states (anamorphs) of lichenized fungi have received little detailed attention from taxonomists. 

Several recent monographic studies have shown that anamorphs can provide useful characters at 

various hierarchial levels of classification and the reader is referred to Vobis (1980) and Vobis & 

Hawksworth (1981) for further background information on the conidial states of lichens. 

Within the genus Micarea there is a diverse array of anamorphic forms, possibly unrivalled by 

any other genus of lichens, except perhaps for some of the genera in the Asterothyriaceae 

(Vezda, 1979) . Information gained from the study of anamorphs has proved invaluable to me for 

the delimitation of species in Micarea; indeed, several species frequently occur without 

apothecia but with numerous pycnidia, such that a detailed knowledge of the latter is often 

essential for their identification (see 'key to species without apothecia'). 

Conidiomata 

The conidiomata are usually pycnidial, and are globose, ovoid, doliiform, or ceriberiform in 

shape. They may be immersed (or partly so) within the thallus or substratum, sessile, or borne 

on stalks (pycnidiophores). When stalked, the pycnidia are usually ± doliiform and the 

stalk-tissue is comprised of loosely interwoven hyphae bound by a gel matrix which is often 

pigmented. In addition, the 'stalk-part' often includes effete pycnidia (Fig. 35B-D). The stalks 

are sometimes branched due to the simultaneous development of two (or more) pycnidia at the


Table 1 Conidial states (anamorphs) found in European species of Micarea. 

Conidium - type Stalked 

Micarea micro- meso- macro- pycnidia 

adnata 

alabastrites 

anterior 

assimilata 

bauschiana 

botryoides 

cinerea 

contexta 

crassipes 

curvata 

denigrata 

elachista 

eximia 

globulosella 

hedlundii 

incrassata 

intrusa 

leprosula 

lignaria 

var. endoleuca 

lithinella 

lutulata 

melaena 

melaenida 

melaeniza + + 

melanobola 

misella + + + 

muhrii 

myriocarpa 

nigella 

nitschkeana 

olivacea 

osloensis 

peliocarpa 

prasina 

pycnidiophora + + 

rhabdogena 

stipitata + + 

subleprosula 

subnigrata 

subviolascens 

sylvicola 

synotheoides 

ternaria 

tuberculata 

turfosa + 

Total (species) 25 28 

apex of stalk tissue (Fig. 35F) or at the apex of an old pycnidium (Fig. 35D) . Stalked pycnidia are 

apparently rare in other genera of crustose lichens (or related fungi). However, I have seen 

branched pycnidiophores associated with lignicolous Chaenothecopsis spp. growing on the 

moribund thaUi and ascocarps of Chaenotheca spp. (several collections in E). 

9 

9


Fig. 35 Micarea botryoides, conidiomata: diagram showing the various means of 'stalk' formation and 

branching (see text for further details). Scale = 100 /xm. 

The pycnidia of species which consistently lack pigment in their apothecia (e.g. M. adnata, M. 

alabastrites, M. pycnidiophora, and M. stipitata) correspondingly lack pigment in their wall 

tissues. In species with coloured apothecia the pycnidial walls are usually (at least in part) 

coloured also. For a given species the pigment involved is usually that found in the hymenium (or 

epithecium) of the apothecia, but in a few cases (e.g. M. lutulata) it is the pigment found in the 

hypothecium which is involved. The link between apothecial and pycnidial pigmentation helps 

distinguish pycnidia belonging to a Micarea from any intermixed pycnidia belonging to a 

non-lichenized or parasitic fungus. It further helps to distinguish the pycnidia belonging to the 

different species of Micarea in mixed populations; for example, the stalked pycnidia of M. 

misella (walls oUvaceous, K+ violet) are often found intermixed with those of other species such 

as M. anterior (walls reddish brown, K— ) and M. melaeniza (walls dark olivaceous, K-). 

When a pycnidium is deeply immersed in the thallus it is often only the uppermost part around 

the ostiole which is coloured, and likewise, if the pycnidium is semi-immersed it is often only the 

upper, exposed part which is coloured. With the exception of the species that are characteristically 

devoid of pigment and some forms of other species in deep shade, the walls of sessile or 

stalked pycnidia are coloured more or less throughout. 

As mentioned above, the conidiomata of Micarea species are pycnidial, but there is one 

exception to this. M. adnata has two anamorphic states: one in which the conidia (mesoconidia) 

are produced internally in immersed pycnidia; and another where the conidia (macroconidia) 

are borne externally on cushion-like sporodochia which resemble small apothecia. This latter 

state could be considered to be a hyphomycetous anamorph but comparative ontogenetic 

studies are required to investigate the theoretical possibility that these sporodochia have 

evolved from pycnidial conidiomata by an exertion of the 'hymenium', and a thickening of the 

subtending wall-tissue to form a supporting cushion (Fig. 36). Unfortunately this hypothesis is 

not supported by intermediate forms in other species, although the pycnidia (with filiform or 

curved-hamate macroconidia) of, for example, M. cinerea and M. peliocarpa often have widely


Fig. 36 Suggested pathway for evolution of a sporodochium (D, as in M. adnata) from a pycnidium (A). 

The step A^B can be seen in the macroconidial conidiomata of e.g. M. cinerea and M. peliocarpa, and 

the mesoconidial conidiomata of e.g. M. denigrata and M. sylvicola. No structure equivalent to 'C is 

known from Micarea. 

gaping ostioles which eventually expose the 'hymenium' . The 

often dubious distinctions 

between pycnidia, acervuli, and other types of conidiomata are discussed by Nag Raj (1981). 

Although the sporodochial anamorph of M. adnata has unusually large, oblong-elHpsoid conidia 

and the longest conidiogenous cells known in the genus, there seems to be no fundamental 

difference in these features from those in other Micarea anamorphs, and the mode of 

conidiogenesis is apparently the same in all cases (see below). 

Conidia 

Vobis & Hawksworth (1981) estimate that perhaps as many as 8000 species of Uchenized fungi 

have conidial anamorphs, and that in most cases a given species has only one conidium type. 

However, it is becoming increasingly evident that a large number of lichens (especially crustose 

lichens) have two conidium types, although many of the examples have not yet been notified in 

the literature. Some examples from my own studies include Anisomeridium biforme, A. 

juistense, Catillaria globulosa, Lecania cyrtellina, Lecanora quercicola, and Opegrapha niveoatra. 

Hedlund (1895) found M. denigrata and M. prasina to each have two conidium types and 

during the present study 16 of the 45 European species of Micarea have been found to have two 

conidium types. Even more surprising has been the discovery that three species {M. denigrata, 

M. nitschkeana, and M. lignaria) each have three conidium types (Figs 42, 45), and to my 

knowledge these are the first reported instances of fungi (whether lichenized or not) with more 

than two pycnidial (coelomycetous) anamorphs. There are, however, a few species of sooty


mould (e.g. in the Metacapnodiaceae) which have three hyphomycetous anamorphs (Hughes 

1972, 1976). 

The main distinguishing features of the three conidium types found in Micarea are as follows 

(see Figs 37-52 for example): Microconidia, narrowly cylindrical or narrowly fusiform, aseptate, 

eguttulate and not constricted in the middle, mostly in the range 3-8x0-5-1 /Am; produced in 

small immersed or ± sessile pycnidia, mostly c. 20-60 /xm diam. Mesoconidia, variable in shape, 

e.g. cylindrical, oblong, ovoid-oblong or obovoid-oblong, aseptate, often biguttulate and (or) 

slightly constricted in the middle; produced in small to large, immersed, sessile or stalked 

pycnidia. All conidia found in stalked pycnidia are included here. Macroconidia, mostly fihform 

or curved and often septate, or heUcoid and septate (M. subnigrata (p. 183)); produced in 

medium-sized to large (often up to 200 jxm diam or more) pycnidia. Also included here are the 

large, aseptate, eguttulate, oblong-ellipsoid conidia produced by the sporodochia of M. adnata. 

In all cases no single pycnidium has been found to contain more than one conidium type. A 

few specimens of M. denigrata and M. nitschkeana have been found with all three anamorphs on 

the same thallus; but such occurrences are rare and it is more usual to find just one or two of 

them. A few species (e.g. M. assimilata, M. crassipes, and M. incrassata appear to have only one 

anamorphic state, with conidia somewhat intermediate between microconidia and mesoconidia 

as defined above; thus the assignment of their conidia to one or other of these types (Table 1) 

must be considered tentative. The distinction between the three conidium types (especially 

micro- and mesoconidia) is most obvious when two or three of them occur on the same thallus. 

The use of the terms 'micro-', 'meso-' and 'macroconidia' are here apphed solely to the conidium 

types found in Micarea - they may not necessarily be analogous to their use in other lichenized or 

non-lichenized fungi. Table 1 indicates the conidium type(s) known for each of the European 

species of Micarea, and from this a numerical summary of the various known combinations is as 

follows: 

Combination of conidium types Number of species 

(anamorphs) (holomorphs) 

Micro- -I- meso- -I- macro- 3 

Micro- + meso- 10 

Micro- -I- macro- 5 

Meso- -I- macro- 1 

Micro- only 6 

Meso- only 

Macro- only 

14 

Anamorph(s) unknown 6 

The role of each of the conidium types is as yet unknown. Vobis (1977) obtained successful 

germination and subsequent growth of mycelium from the macroconidia of Lecanactis abietina, 

but was unable to germinate the microconidia of the same species. His results suggest that the 

former may well act as asexual propagules, and the latter as spermatia in sexual reproduction. It 

seems most likely to me that the microconidia oi Micarea species are spermatia. As to whether or 

not they are essential components of the sexual reproductive process is much less certain; 21 of 

the treated species are not known to produce microconidia. 

The role of conidia as asexual propagules is still a matter of much debate and speculation 

among lichenologists, but such a role (at least in certain cases) can be inferred from the fact that a 

few crustose species (e.g. Lecanactis subabietina and an undescribed Bacidia) are not known to 

have apothecia or vegetative diaspores (i.e. soredia, isidia, etc.) but always have numerous 

pycnidia. To such examples can be added the numerous crustose lichens whose apothecia are 

known, but which commonly occur as sterile populations with numerous pycnidia, e.g. 

Anisomeridium juistense (macroconidial state), Arthonia phaeobaea, A. spadicea, Bacidia 

arnoldiana, B. carneoglauca, B. trachona, Cliostomum graniforme, C. griffithii, Lecanactis 

abietina, Lecidea erratica, Opegrapha niveoatra, O. vermicellifera, and O. vulgata. This last 

group also includes Micarea botryaides, M. denigrata, M. misella, M. pycnidiophora, and M. 

stipitata, all of which commonly occur with abundant mesoconidia-containing pycnidia and few 

(if any) mature apothecia.


Fig. 37 M. adnata (E - holotype), macroconidia and conidiogenous cells. Scale = 10 />tm.


00 B 

Fig. 38 A, M. botryaides (Coppins 8429, E), mesoconidia and conidiogenous cells. B, M. melaeniza (S - 

holotype), mesoconidia. C-D, M. anterior Sweden, Angermanland, Langsele, 1892, Hedlund, S); C, 

mesoconidia; D, microconidia. Scale = 10 yam.


s 

o 

o 

c 

o 

c 

o 

-o cCO 

u 

6 

I 

ei)

\r\ r\ 

[J^\J 


r\ 

\J 

Fig. 40 A-B, M. contexta (S - lectotype); A, mesoconidia; B, microconidia. C, M. eximia (Malme Lich. 

suec. exs. 26, C), mesoconidia. D, M. bauschiana {Coppins 4111, E), microconidia and conidiogenous 

cells. Scale = lOjum.


n 

1/ 

[\ f] n 

Fig. 41 A-B, M. incrassata, microconidia; A, (S - holotype); B, (Kerguelen, 1875, Eaton, E). C, M. 

crassipes (Vezda Lich. Sel. 11, BM), (?)mesoconidia and conidiogenous cells. Scale = 10 /tm. 

B


Fig. 42 M. denigrata (Coppins 1888, E), conidia with conidiogenous cells; A, macroconidia; B, mesoconi- 

dia; C, microconidia. Scale = 10 /Am.


Fig. 43 A-B, M. globulosella (Czechoslovakia, Slovakia, Vysoke Tatry, 1879, Lojka, BM); A, mesoconidia; 

B, microconidia. C, M. globulosella (S - lectotype), mesoconidia. D-E, M. synotheoides (Coppins 

2942, E); D, mesoconidia; E, microconidia. Scale = 10 /jun.


Fig. 44 A, M. hedlundii (UPS - holotype), mesoconidia and conidiogenous cells. B, M. lutulata (Wales, 

Pembroke, Tycanol, 1980, James, BM), mesoconidia and conidiogenous cells. Scale = 10 )Ltm.


Fig. 45 M. lignaria. A, macroconidia (Coppins 4658, E). 

microconidia (Coppins 8952, E). Scale = 10 ixm. 

B 

B, mesoconidia (Coppins 8436, E). C,


\j \l 

Fig. 46 A, M. melaena {Coppins 6041, E), macroconidia. B-C, M. melanobola (H-NYL 2164 

lectotype); B, mesoconidia; C, microconidia. Scale = 10/i,m. 

B


B 

OOQog 

Fig. 47 A, M. olivacea (BM - holotype), mesoconidia and conidiogenous cells. B, M. nigelta (E - 

holotype), mesoconidia and conidiogenous cells. C, M. myriocarpa (Coppins 8939, E), mesoconidia and 

conidiogenous cells. Scale = 10 /xm.


Fig. 48 M. peliocarpa (England, New Forest, Great Wood, Bramble Hill Walk. 1970, Coppins et al., E). 

A, macroconidia and conidiogenous cells. B, microconidia and conidiogenous cells. Scale = 10 jxm.


Fig. 49 A-B, M. prasina (GZU - holotype of M. polytrichi); A, mesoconidia; B, microconidia. C, M. 

prasina (Coppins 8009, E), mesoconidia. D, M. prasina {Coppins 2835, E), microconidia. Scale — 10 

fim. 

{\ 

^ 

B


Fig. 50 M. subnigrata (Coppins 8417, E). A, macroconidia and conidiogenous cells. B, microconidia and 

conidiogenous cells. Scale = 10 /u,m. 

B


Fig. 51 A, M. sylvicola {Muhr 2579, E), mesoconidia and conidiogenous cells. B, M. tuberculata (O - 

lectotype), mesoconidia and conidiogenous cells. Scale = 10 /x,m. 

B


Fig. 52 A-B, M. ternaria, mesoconidia; A, (H-NYL 18682 - holotype); B, {Thomson 9188, DUKE). C, 

M. aff. ternaria (Fair Isle, 1976, Duncan, BM), mesoconidia. Scale = 10 fjun. 

B

84 


In the case of M. denigrata it is a general rule (but with exceptions) that when on worked wood 

(fence-posts, garden furniture, window frames, etc.) its thallus has numerous pycnidia (with 

mesoconidia) and often few (if any) mature apothecia. On natural substrata (e.g. fallen tree 

trunks in old woodlands) pycnidia with microconidia or macroconidia are more prevalent, 

although they are usually relatively fewer in number and associated with numerous apothecia. It 

seems highly likely that the success of M. denigrata as a primary coloniser of newly available 

substrata is largely due to the successful role of its mesoconidia as asexual propagules. Its 

microconidia probably function as spermatia, but the function of the macroconidia is less 

obvious, although I suspect that they, like the mesoconidia, act as asexual diaspores (perhaps in 

a different way). 

Another observation pertinent to this discussion is that I have frequently observed the mesoand 

macroconidia of Micarea species being extruded through the ostioles of the pycnidia as 

white mucilaginous blobs; such phenomena are usually seen after periods of wet weather. It is 

tempting to suggest that these extrusions facilitate dispersal beyond the limits of the parent 

thallus by the action of rain or passing arthropods and mollusca. Dispersal by invertebrates may 

be of especial importance for species such as M. botryoides, which occur in sheltered situations 

rarely subjected to rain-wash. I have only rarely observed microconidia being extruded as white 

blobs, thus suggesting that there is no need for them to be dispersed beyond the limits of the 

parent thallus. Indeed, if microconidia do function as spermatia, and if the mycobiont of the 

Micarea is homothallic, there would be no requirement for them to be dispersed more than a few 

millimetres. 

Conidiogenous cells 

The conidiogenous cells of Micarea species are always phialidic, although in many cases the 

phialides are seen to undergo 'percurrent proliferation' (Figs 42B, 44B). The conidiogenous 

cells belong to Types I or II of Vobis & Hawksworth (1981) and arise from the inner wall of the 

pycnidium (or on the outer surface of the sporodochium in M. adnata). Their subtending cells 

are never sufficiently regular in shape for them to be termed 'conidiophores'. In several species 

whose pycnidial walls are intensely pigmented the bases of the conidiogenous cells are often 

similarly pigmented (Figs 47B, 51B). The nearest approach to the differentiation of wall-tissue 

from a conidiogenous layer is found in the thick-walled pycnidia of M. elachista. 

There is much variety in the shape of conidigenous cells (e.g. ampuUiform, doliiform, 

lageniform, cyUndrical) and there is sometimes much variation within a single pycnidium 

(Figs 42B, 44A). Conidiogenous cells with long cyhndrical necks often have swollen bases (Figs 

38A, 47B, 5 IB). Although critical observations and measurements have not been made for all 

species, the size and shape of conidiogenous cells have rarely been found to be useful characters 

for the separation of closely similar species. However, one exceptional case is that of saxicolous 

forms of M. olivacea (Fig. 47A) versus M. tuberculata (Fig. 5 IB), in which the conidiogenous 

cells of the latter are much longer. 

Chemistry 

The discussions below are presented under two sub-headings. The first deals with 'lichen 

substances', which are readily extractable in acetone and identifiable by thin-layer chromatography 

(t.l.c; see 'Methods'). The second part deals with pigments that cannot be analysed in this 

way; their chemical nature is at present unknown and they can only be characterised by their 

colour in water and subsequent reactions with reagents such as K and HNO3. 

Lichen substances 

Prior to the present studies there is little evidence in the literature to suggest that species of 

Micarea contain any lichen substances. However, there are a few hints given in some early 

descriptions: for example, Leighton (1879: 362) gives 'K+ yellow, C+ orange-red' reactions for 

Lecidea milliaria [Micarea lignaria], which probably relate to his specimens of the var. 

endoleuca. The only reported t.l.c. analysis of a Micarea appears to be that of Huneck &


Table 2 Chemical content and corresponding spot test reactions of European species of Micarea known to 

contain lichen substances. +, present; ±, presence variable and sometimes absent; ?, present occa- 

sionally as trace amounts or contaminant; R, red; Y, yellow; fY, faint yellow; O, persistent orange; ( ) 

reactions not obtainable in many collections; *, including 3 or 4 accessory substances.


6 - 

5h 

4 

3 

2 

1 

r m^ 

B 

C 

F F 

D D 

Fig. 53 Diagram of chromatogram in solvent system TDA. 

^ 

K 

J 

(22) (^ 

8 

TDA 

1, Control (Parmelia acetabulum plus Cladonia subcervicornis). 2, M. peliocarpa. 3, M. subleprosula. 4, 

M. lignaria. 5, M. leprosula. 6, M. lignaria var. endoleuca. 7, M. prasinas. lat. 8, M. prasinas. str. 9, M. 

prasina s. lat. A, atranorin, B, norstictic acid. C, fumarprotocetraric acid. D, gyrophoric acid. E, 

alectorialic acid, and accessory substances (E\ E^). F, argopsin. G and H, xanthones. J, prasina 

unknown A. K, prasina unknown B. L, prasina unknown C. 

7 r 

6 

5 

4 

3 

2 

1 

B 

(Si 

D


most widely occurring compounds among lichen genera. It has been found in 11 Micarea species, 

although its presence in M. curvata has only been inferred from spot tests, the solitary specimen 

(holotype) being too small for t.l.c. analysis. In addition to these occurrences, gyrophoric acid is 

frequently detected in trace amounts in chromatograms of other taxa, especially M. prasina s. 

ampl. The fact that Micarea species often grow intermixed with other lichens, including several 

that contain gyrophoric acid (e.g. Lecidea icmalea and L. granulosa agg.) raises the question as 

to whether these trace amounts represent its presence as an accessory substance or as a 

contaminant. This is difficult to answer. Gyrophoric acid is certainly found in the M. prasina 

complex, as it has been detected in large amounts in the type material of Lecidea levicula Nyl. 

from Cuba. (Further studies are required to establish the taxonomic status of L. levicula.) 

Well-developed specimens of M. denigrata, M. nitschkeana, and M. melaena have an areolate 

thallus containing readily detectable quantities of gyrophoric acid. However, the thallus of these 

species is sometimes scurfy-granular and blackish due to disruption by invading dematiaceous 

fungi and non-lichenized algae, and gyrophoric acid is produced in very low amounts or is 

apparently absent altogether (not detectable by t.l.c). 

The most surprising result of the chemical studies in Micarea was the discovery of three 

unknown, but very distinctive, compounds in M. prasina. The three compounds can be 

characterised thus (UV at 254 m/x): 

prasina unknown A. TDA 5: HEF 6, UV-h blue-white. After H2SO4 and charring: UV+ dull 

orange-red, dull orange in daylight. 

prasina unknown B. TDA 5: HEF 6, UV+ blue (less bright than 'unknown A'). After H2SO4 

and charring: UV-f- vivid citrine-yellow, yellow (without orange tinge) in daylight. 

prasina unknown C. TDA 4-5: HEF 6-7 (slightly higher than 'unknowns A and B'), UV-I- 

grey or pale mauve (TDA) or ± colourless (HEF). After H2SO4 and charring: UV-I- violet-blue, 

± colourless in daylight but turning a pale pinkish-lilac after several weeks. 

In all the numerous European and North American specimens of M. prasina s. ampl. 

examined so far, these compounds have never been found in combination, and M. prasina exists 

as three distinctive chemical races (see taxonomic account of M. prasina for further discussion). 

None of the three compounds can be identified in the tabulations of Culberson (1972), and they 

have never been encountered in other genera during the numerous and diverse investigations by 

Mr P. W. James (pers. comm.). Samples of specimens containing 'unknowns A and B' are 

currently being studied by Dr J. A. Elix. 

The identification of hchen substances (even if only by spot tests) is essential for distinguishing 

sterile specimens of M. leprosula from M. subleprosula, and for separating the two varieties of 

M. lignaria; it is also of great value in the routine identification of many other species (see 'Keys 

to species'). The three European races of M. prasina are not distinguishable by spot tests, but are 

readily identified by t.l.c. 

Pigments 

The nature of pigments and their location (especially within apothecia) are of prime importance 

in the delimitation of species within many lichen genera, particularly those included in the 

Lecideaceae, and Micarea is no exception. Little is known of these acetone insoluble pigments 

and a detailed knowledge of their structure and biogenesis would be invaluable for the 

evaluation of their taxonomic significance. 

On the basis of the colours seen in water mounts and the colour changes brought about by the 

application of a strong alkali (KOH) and a strong acid (HNO3) at least eight pigments can be 

recognized in Micarea. A very provisional summary of these pigments is given below: 

Pigment A. Green or aeruginose, K- or -I- green intensifying, HNO3+ red; in various tissues (according 

to species) of M. assimilata, M. bauschiana, M. cinerea, M. intrusa, M. lignaria, M. melaena, M. 

peliocarpa, M. sylvicola, and M. tuberculata. 

Pigment B. Purple, K+ green, HNO3+ purple-red; mostly in the hypothecium of M. assimilata, M. 

contexta, M. crassipes, M. eximia, M. melaena, M. nigella, and M. sylvicola (rare forms). 

Pigment C. Purple, K+ purple intensifying (sometimes partly dissolving into solution); occasionally in

. 


the epithecium (M. melaenida and forms of M. crassipes and M. melaend) but mainly in the hypothecium 

{M. assimilata, M. crassipes, M. melaena, M. melaenida, and M. subviolascens) 

Pigment D. Olivaceous, K+ violet (also C+ violet), HNO3+ red; in various tissues (but rarely in the 

hypothecium, and then in low concentrations) of M. denigrata, M. elachista (mostly in pycnidia), M. 

globulosella, M. hedlundii, M. melanobola, M. misella, M. nitschkeana, M. prasina, M. subviolascens, and 

M. synotheoides 

Pigment E. Fuscous brown, K+ dissolving into solution, HNO3—; in the epithecia of M. elachista SiXxdM. 

rhabdogena. 

Pigment F. Brown (sometimes slightly tinged reddish), K- (not dissolving), HNO3- or + orangebrown; 

in various tissues (according to species) of A/, botryoides, M. curvata, M. incrassata, M. lutulata, M. 

muhrii, M. myriocarpa, M. osloensis, M. subnigrata, andM. turfosa. It is quite possible that more than one 

pigment is involved here. 

Pigment G. Dilute yellowish, K+ purple (oily droplets), HNO3- ; in goniocysts and sometimes the lower 

hymenium and hypothecium of M. hedlundii. 

Pigment H. Pale yellowish orange, K+ purple, HNO3— ; in cytoplasm of some ascogenous hyphae, asci, 

and spores of M. intrusa. This is possibly similar (or identical) to pigment G. 

Pigment complexes involving mixtures of pigments A, B, and C are often found (especially in 

the hypothecium) in M. assimilata, M. contexta, M. crassipes, M. melaena, M. subviolascens, 

and M. sylvicola; see the individual species accounts for further details. The existence of such 

complexes suggests that pigments A, B, and C are closely related chemically. 

Pigment A is probably identical to that found (usually in the epithecium) in a large number of 

lichens, especially in Ledêfl 5. lat., Catillarias. lat., Bacidias. lat. and Lecanoras. lat. Pigment 

C is probably of rare occurrence outside Micarea, but I know of it in the epithecium of Bacidia 

beckhausii and in several species of Pertusaria (e.g. P. hymenea and P. oculata). I have not 

encountered pigment E in any other lichens, but it should be compared with the pigment(s) 

responsible for the ionomidotic reaction found in several genera of non-lichenized discomycetes 

(Korf , 1973). Pigment F is probably of wide occurrence, but there may be several pigments that 

impart a brown or 'melanized' appearance in tissues. 

The dark violet-blue, K-h aeruginose granules occasionally seen in the hymenium of M. 

contexta and M. lignaria (and also the apothecia of Lecidea hypnorum and Dactylospora 

lobariella), is probably related to (if not the same as) pigment B, and it may be the same as the 

epithecial pigment found in Bacidia absistens, Mycoblastus fucatus , and Schaereria tenebrosa. 

Ecology 

No detailed ecological studies have been made in connection with this primarily taxonomic study 

of Micarea, and much of the discussion given below is based on floristic notes and casual field 

observations. For most amateur and professional lichenologists alike, the species oi Micarea are 

little known, poorly understood, and much overlooked in the field. For this reason the following 

account is more of a guide to collectors, rather than an ecological dissertation. Additional 

ecological notes are also included in the taxonomic accounts of each of the Micarea species 

treated. Syntaxonomic nomenclature follows James ef a/. (1977). 

General habitats 

Micarea species occur in a wide range of habitats but are confined to substrata with a low pH 

(below c. pH 5) and generally avoid nutrient enriched situations. Thus, they are absent from 

limestone rocks, tops of bird-perching stones, seashores, and basic bark. There are a few partial 

exceptions, for example M. denigrata can occur on wooden fencing and other timber in 

farmyards, M. prasina can occur on soil and debris among rocks in the upper seashore, and M. 

lignaria has been found on exposed limestone growing over mats or cushions of moribund 

bryophytes from which free calcium ions have presumably been leached out. The bark of Acer 

and Ulmus normally has a pH which is too high for species of Micarea (with the exception of M. 

prasina in some situations), but in areas heavily polluted by sulphur dioxide and its derivatives 

(e.g. West Yorkshire Conurbation), bark pH can be substantially lowered (Gilbert, 1970), thus 

providing a suitable substratum for species such as M. melaena (q.v.) and M. botryoides (q.v.). 

.


The principal habitats (I-XI) in which Micarea spp. are found are given below; each is 

provided with a list of the species which can be expected to occur in it (allowing for climatic and 

phytogeographical variations). Species given in square brackets [ ] are rarely and unusually 

found in the given habitat, and those prefixed with a dagger (t) are not known in this habitat 

from the British Isles. 

I. 

Sheltered, dry underhangs, usually in valley woodland or narrow ravines, including dry undersides 

of up-ended tree root systems in woodland; growing on rock, loose stones, consolidated soil, and 

exposed roots. 

M. bauschiana M. myriocarpa 

M. botryoides M. sylvicola 

M. lignaria M. tuberculata 

M. lutulata 

II. Exposed parts of sheltered rocks, usually in woodland; more frequently wetted than I; growing 

directly on rock. 

M. botryoides M. sylvicola 

tM. curvata [M. cinerea] 

M. lignaria s. lat. [M. denigrataj 

M. lithinella '\IM. muhrii] 

M. melaena [M. nitschkeana]^ 

M. olivacea [M. prasina] 

M. peliocarpa 

a, stones amongst Calluna in heathland. 

III. Over bryophytes on rocks, boulders, old stumps, or fallen trees in woodland at low altitudes (500 m 

in the British Isles). 

M. adnata M. lignaria s. lat. 

M. botryoides M. melaena 

M. cinerea M. peliocarpa 

M. leprosula M. prasina 

[M. stipitata] 

IV. Exposed, hard siliceous rocks; growing directly on rock. 

M. intrusa tM. subviolascens 

M. lignaria M. aff . ternaria^ 

M. subnigrata 

b, coastal districts only. 

V. Over bryophytes or peaty debris on exposed turf or on, or amongst, rocks and boulders in open 

situations in upland, montane, or 'arctic' districts. 

M. assimilata M. peliocarpa 

tM. crassipes M. subleprosula 

M. incrassata tM. ternaria 

M. leprosula M. turfosa 

M. lignaria s. lat. [M. cinerea] 

M. melaena 

VI. Over bryophytes or plant debris on the ground in old dunes, disused lead and zinc mines, sea-cliffs, 

or by woodland tracks. 

M. botryoides M. prasina 

M. leprosula [M. denigrataj 

M. lignaria s. str. flM. misella] 

M. peliocarpa 

VII. On bare mineral soil at low altitudes. 

tM. melaenida [M. lignaria] 

tM. osloensis [M. prasina] 

[M. leprosula] 

VIII. Corticate trunks of healthy trees; usually in woodland; sometimes overgrowing bryophytes (*). 

*M. alabastrites *M. pycnidiophora 

*M. cinerea *M. stipitata


fM. elachista *M. synotheoides 

tM. globulosella [M. botryoides] 

M. melaena [M. denigrata] 

tM. melanobola [M. leprosula] 

*M. peliocarpa [M. lignaria] 

*M. prasina [M. nitschkeana] 

IX. On attached twigs of trees or large shrubs, or thin stems of small shrubs; in woodland, heathland, or 

scrub. 

tM. cinerea M. peliocarpa 

M. lignaria s. str. M. prasina 

M. nitschkeana [M. denigrata] 

X. On timber (i.e. worked wood), e.g. fencing, garden furniture, old window frames, and shingles. 

M. cinerea M. nitschkeana 

M. denigrata M. peliocarpa 

tM. elachista [M. globulosella] 

M. lignaria s. str. [M. leprosula] 

M. melaena [M. sylvicola] 

M. misella 

XI. Directly on lignum of old stumps and decorticate trunks. 

M. adnata M. misella 

M. alabastrites tM. muhrii 

tM. anterior M. nigella 

M. cinerea M. nitschkeana 

tM. contexta M. olivacea 

M. denigrata M. peliocarpa 

tM. elachista M. prasina 

tM. eximia tM. rhabdogena 

tM. hedlundii /M. botryoides] 

M. lignaria s. lat. [M. leprosula] 

M. melaena [M. myriocarpa] 

tM. melaeniza [M. synotheoides] 

From the above lists it will be seen that several species (e.g. M. lignaria, M. melaena, M. 

peliocarpa, and M. prasina) inhabit a very wide range of habitats and substrate. At the other 

extreme there are many species which are much more restricted e.g. M. tuberculata (I), M. 

subnigrata (IV), M. assimilata (V), M. melaenida (VIII), M. pycnidiophora (VIII), and M. 

anterior, M. contexta, M. eximia, and M. melaeniza (XI). 

In most of the communities in which they occur Micarea species are usually of minor 

importance with regard to cover values. However, there are some exceptions such as the 

Micareetum sylvicolae association of underhangs and exposed tree root-system (see below), the 

community dominated by M. prasina on trunks in dense conifer plantations, some lignicolous 

assemblages on fallen trunks, old stumps, and worked wood. Micarea nitschkeana on Calluna 

twigs and litter in some lowland heaths, M. melaena on sandy or peaty soil in some heathlands 

and moorlands, and M. lignaria on the ground in some old lead mine workings. 

Specific habitats 

Deciduous (broad-leaved) woodland 

In Britain the genus JWicarea is best represented in terms of number of species per site in the 

mature, ± natural woodland (both deciduous and coniferous) on acid soils, in areas with a high 

annual rainfall (at least 1000 mm distributed over at least 160 'wet days'; see Coppins, 1976). The 

best examples of 'Mcarefl-rich' deciduous (broad-leaved) woodlands are found in Wales, the 

English Lake District and the west of Scotland north of the Clyde Estuary. The Micarea species 

found in them on trees and stumps are included in lists III, VII, and XI above, except that M. 

misella, M. nigella, and M. olivacea are primarily species of coniferous woodlands. 

The occurrence of Micarea spp. on bark (or over bryophytes thereon) is favoured by leaching, 

and is further favoured, to the detriment of more basicolous lichens, by the effects of 'acid rain'


(Anon, 1980; Fowler et al., 1982). Acid rain effects, resulting from the pollution emitted from 

the industrial areas of west-central Scotland (Clydeside and Glasgow conurbation), are probably 

the explanation for the prevalence on the trunks of mature trees (e.g. Quercus, Alnus, 

Betula, and Fraxinus) of such species as M. alabastrites, M. cinerea, M. peliocarpa, M. stipitata, 

and M. synotheoides in the deciduous woodlands of the Cowal Peninsula in southern Argyllshire. 

The communities in which these species occur are probably referable to the Parmelietum 

laevigatae, and in areas subjected to 'acid rain' it appears that these communities have to some 

extent, and in certain situations (especially on Quercus and Fraxinus), replaced the more 

basicolous communities of the Lobarion pulmonariae. 

In the deciduous woodlands of drier parts of the British Isles (annual rainfall of


Saxicolous habitats 

Of the 45 European species of Micarea, 21 have been found growing directly on rock (lists I, 

II, and IV), although the genus is poorly represented in the main saxicolous lichen alliances of 

exposed siliceous rocks (e.g. Lecideion tumidae, Rhizocarpon alpicolae, and Umbilicarion 

cylindricae; list IV). As saxicoles Micarea species are more prevalent in sheltered ravines and 

woodlands. In dry underhangs in rock faces, steep banks, and below overhanging trees, up to 

seven species (Hst I) may be found in the ombrophobous, aerohygrophilous Micareetum 

sylvicolae. The species in this community grow on rock, loose or lodged stones, exposed roots, 

consolidated soil, and encroaching dry mats of bryophytes. Associated lichens from other 

genera may include Coniocybe furfuracea, Enterographa hutchinsae, Melaspilea subarenacea, 

Microcalicium arenarium, Opegrapha gyrocarpa, O. zonata, Porina chlorotica, P. lectissima, 

Psilolechia clavulifera, and P. lucida. This community is usually well defined, but in some 

situations it intergrades with other assemblages of shaded rocks such as the Lecideetum lucidae, 

Coniocybetum fururaceae, Opegraphetum horistico-gyrocarpae, and the Racodietum rupestris. 

A few Micarea species (list II) may be found on the upper sides of stones, boulders, and rocks 

where they are subjected to direct wetting by rain, or by drips from the overlying tree canopy. In 

most cases these are incidental occurrences of species more characteristic of other substrata. 

However, such situations may be the normal habitat of the little known M. curvata and M. 

lithinella. 

In northern and western Britain several Micarea species occur on mossy rocks. The communities 

involved are often difficult to place, some belonging to bryophyte-dominating syntaxa, 

others to normally epiphytic communities (e.g. Parmelietum laevigatae), or to chomophytic 

communities dominated by bryophytes and Cladonia species which, in turn, are often invested 

by a gelatinous algal scum. 

Terricolous habitats 

Areas of lowland heathlands where the peaty or sandy soil has been laid bare of tall vegetation 

by erosion, disturbance, or burning are colonised by several species of algae bryophytes and 

lichens. The lichens involved often include species such as Lecidea icmalea, L. uliginosa, L. 

oligotropha (rare in Britain), Baeomyces rufus, B. roseus, and some Cladonia spp. Micarea 

species are not usually involved in such communities, although M. melaena may at times be 

present, occasionally achieving local dominance. In mature Callunetum in the heathlands of 

eastern England and Jylland (Denmark) M. nitschkeana is sometimes encountered in abundance 

on litter, as well as on the thin attached twigs of the Calluna. Acidic soils contaminated by 

heavy metals (especially lead) are often rather bare with an open vegetation, and Micarea 

species (especially M. lignaria) may occur in quantity growing over moribund bryophytes and 

plant debris. Soil, plant debris, and moribund bryophytes amongst coastal rocks are often 

colonized by M. prasina (q.v.). M. melaenida (q.v.) is found exclusively on consohdated fine 

grained (argillaceous) mineral soils. The terricolous species encountered in upland, montane, or 

arctic regime are given in list V; the reader is referred to the individual species accounts for 

further details. 

Man-made substrata 

A detailed treatment of the lichens of man-made substrata is given by Brightman & Seaward 

(1977). Several Micarea species occur on worked wood (list X), but M. denigrata {q. v. ) is by far 

the most successful species. M. denigrata has also been found on pieces of hardboard lying in a 

dune slack at Tentsmuir in Fife. Old sackcloth and other fabrics lying on the ground in old 

lead-mine workings are frequently colonised by lichens, including M. lignaria. A more surpris- 

ing find was that of M. nitschkeana, growing with Scoliciosporum umbrinum, on a small plastic 

carton in a heathland on the Isle of Wight. Further field studies will undoubtedly extend this list 

of artificial substrata. Micarea denigrata and M. prasina are rapid colonisers of newly available 

substrata, and have been collected respectively on dead culms of Cladium mariscus and 

Phragmites australis in natural habitats. Both of these reeds are sometimes used as thatch, and a 

close inspection of thatched roofs ought to reveal the presence of Micarea species.


Habitats in lowland pastoral areas 

The lowlands of eastern England, the English south midlands and parts of eastern Scotland 

contain few 'natural' habitats suitable for Micarea species. In addition, these same areas suffer, 

at least to some extent, from the effects of incoming pollution (especially sulphur dioxide and its 

derivatives) and indigenous pollution in the form of agricultural biocides and fertilisers. 

Nevertheless, the diligent collector will usually discover suitable niches where Micarea species 

are to be found. The most commonly encountered member of the genus in lowland agricultural 

and suburban districts is M. denigrata which can grow on worked wood in a wide range of 

situations, and on old tree stumps in hedgerows. M. prasina can be found on trees and shrubs in 

small woodlands planted as fox coverts or for the rearing of pheasants. It may also be found on 

sheltered stems of shrubs in waste ground, marginal land, or gardens, where M. nitschkeana may 

also occur. 

A major refuge for wildlife (including lichens) in lowland Britain is provided by churchyards 

(Anon, 1973, 1978). Although records of Micarea species are rather few, M. denigrata can often 

be found on timber and, more rarely, on siliceous memorial stones; and M. lignaria and M. 

peliocarpa have been recorded on the siliceous stone-work of old walls and memorials. 

Distribution 

Britain 

Maps 1-28 present the distributions of all the Micarea species occurring in the British Isles, with 

exception of M. lithinella which is known from just one 10 km grid square (44/86). It is seen from 

these maps that records for most species are concentrated in the western and/or northern 

districts. This is partly due to the fact that suitable habitats are most numerous in these areas, 

and it is probable that several species (e.g. M. bauschiana, M. lignaria, M. melaena, M. 

peliocarpa, and M. sylvicola) would be more evenly distributed if there were more available 

habitats in the lowland areas of the south and east. It is impossible to discuss in isolation the 

effects of climate or substrate availability on plant distributions. Substrate availability is largely 

dependent on topography and geology, but the physical and chemical nature of an existing 

substratum can be much influenced by the prevailing climate. For example, the bark of trunks of 

mature Quercus in high rainfall districts tends to be more leached, soft, and friable (i.e. more 

suitable for Micarea spp.) than in low rainfall districts. As a second example the species of the 

Micareetum sylvicolae (see p. 92) are ombrophobous (avoid frequent direct wetting), but are 

also aerohygrophilous (require ± constant high humidity). In western Britain the high rainfall, 

high incidence of cloud cover (i.e. low duration of possible sunshine), and incoming moist 

air-stream from the Atlantic on the prevailing westerly winds result in high levels of relative 

humidity (and low levels of saturation deficit) more or less throughout the year (see Climato- 

logical Atlas of the British Isles, London: HMSO (1952)). Under such conditions a well 

developed Micareetum sylvicolae can be found on underhangs in a wide range of lowland 

habitats. Eastern districts generally have a lower rainfall and lower incidence of cloud cover 

resulting in lower levels of relative humidity, such that the Micareetum sylvicolae tends to be 

more confined to narrow river valleys with a long history of continuous tree cover. A western 

bias to the present day British distribution of many lichens is often due to the fact that the eastern 

side of the country is generally that most affected by industrial and urban development, 

intensive agricultural practices, and the resultant forms of air pollution (Hawksworth, Coppins 

& Rose, 1974; Coppins, 1976). 

A climatically determined western distribution, attributable to the General Western Group of 

Coppins (1976), is shown by Micarea adnata, M. alabastrites, M. cinerea, M. lignaria var. 

species have a eu- Atlantic or sub-Atlantic 

endoleuca, M. stipitata, and M. synotheoides . These 

European distribution, and are confined to areas with an annual rainfall of over 800 m (mostly 

over 1000 mm) distributed over at least 160 wet days ['wet day' = period of 24 h in which 1 mm 

(or more) of rain is recorded]. The genus Micarea is poorly represented in southern (Mediterranean) 

Europe, and it is not surprising that few British species exhibit a marked southern 

tendency. The best example of such a species is M. pycnidiophora, which is mainly confined to


The identification (and subsequent annotation and cataloguing) of a single specimen of 

Micarea (which more often than not is fragmentary or in poor condition) can be very time 

consuming, and often involves lengthy microscopical examination. The unfortunate consequence 

of this is that I have not been able to examine a large number of potentially available 

specimens from several major institutional and private herbaria. 

Despite the above problems and shortcomings, I have tentatively attempted below to assign 

the species of Micarea to some general distributional types. In most cases I have paid little 

attention to purely literature sources of records because I have found that many such reports, 

even those by lichenologists whose works I hold in the highest esteem, can be unreliable in 

respect of current species concepts. 

A. Eu-Atlantic - mainly confined to oceanic areas including west Norway, the British Isles, western 

France, and Macaroenesia. 

M. alabastrites M. stipitata 

M. nigella M. subnigrata 

M. olivacea tM. subviolascens 

*M. prasina (with 'unknown C) M. synotheoides 

*M. pycnidiophora 

*with southern tendency twith affinity to group G 

B. Sub-Atlantic - as above, but also present in high rainfall (annual rainfall of >800 mm) areas of 

central Europe (Alps, Carpathians, etc.). 

M. adnata M. lignaria var. endoleuca 

M. cinerea 

C. Boreal - mainly confined to Fennoscandia, but absent or rare from west Norway. 

M. anterior M. melanobola 

M. contexta M. muhrii 

M. eximia M. osloensis 

M. melaeniza M. rhabdogena 

D. Boreal-continental - present in Fennoscandia and central Europe, but rare or absent from west 

Norway and western Britain. 

M. elachista M. lithinella 

M. hedlundii *M. melaenida 

*but known from western France 

E. Continental - known only from central Europe. 

M. curvata 

F. Montane (arctic-alpine) - at high altitudes in montane regions of Britain and central Europe, but 

sometimes at low altitudes in Fennoscandia. 

M. assimilata M. subleprosula 

M. crassipes M. turfosa 

M. incrassata 

G. Arctic - mainly confined to within the Arctic Circle, but possibly extending southwards along the 

Atlantic coast. 

M. ternaria 

H. Widely Distributed - known from British Isles, Fennoscandia, central Europe, and often elsewhere. 

More records will probably reveal distinct phytogeographical tendencies for some of the 

species included here. 

M. bauschiana M. melaena 

M. botryoides tM. misella 

M. denigrata M. myriocarpa 

XM. globulosella M. nitschkeana 

M. intrusa XM. peliocarpa 

M. leprosula *M. prasina 

M. lignaria var. lignaria 

M. sylvicola 

M. lutulata 

*races containing 'prasina unknowns A and B' 

twith possible affinity to group D 

twith possible affinity to group B 

M. tuberculata


World 

The present study is confined to species of Micarea that occur in Europe. However, some 

extra-European specimens have been examined in connection with nomenclatural matters, and 

others have been examined incidentally. According to current information the genus is best 

represented in Europe, but this may, or may not, be true. Micarea is well represented in North 

America from where I have seen 12 species (all European taxa: M. crassipes, M. denigrata, M. 

globulosella, M. lignaria s. sir., M. melaena, M. misella, M. nitschkeana, M. peliocarpa, M. 

prasina, M. sylvicola, M. ternaria, and M. turfosa); this list will undoubtedly be much extended 

in the near future. From other regions I have seen (but not necessarily critically examined) 

specimens of Micarea from Japan, Borneo, Tasmania, New Zealand, South Africa, South 

America (Brazil), and the Antilles (Cuba). It is likely that species of Micarea are to be found in 

most temperate and boreal regions, as well as in many tropical regions (especially mountainous 

areas). The numerous collections that I have received from New Zealand include several 

undescribed taxa but they also include at least one European species, Micarea peliocarpa. Three 

additional European species known from the southern hemisphere are M. incrassata (Ker- 

guelen), M. lignaria (Brazil), and M. misella (Brazil). 

Micarea Fr. 

The genus Micarea 

Syst. orb.: 256 (1825). - Micarea Fr. emend Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18(3): 27 

(1892). Lectotype: Micarea prasina Fr. (see note below). 

Helocarpon Th. Fr., Lich. arctoi: 178 (1860); Nova Acta R. Soc. Sclent. Upsal. Ill, 3: 278 (1861). Type 

species: Helocarpon crassipes Th. Fr. [= Micarea crassipes (Th. Fr.) Coppins]. 

Stereocauliscum Nyl. in Flora, Jena 48: 211 (1865). Type species: Stereocauliscum gomphillaceum Nyl. 

[= Micarea lignaria (Ach.) Hedl.] 

Micarea sect. Bryophagae Poelt & Dobbeler in Bot. Jb. 96: 337 (1975). Type species: Micarea polytrichi 

Poelt & Dobbeler [= Micarea prasina Fr.]. 

Note. The name Micarea was first validly published in December 1825 by Fries {op. cit.), although it was 

twice mentioned by the same author earlier in the same year. In Sched. crit. lich. siiec. part 3, fasc. 4, page 

21 (pre 7 May) it appeared in a note under the entry for Biatorafuliginea. However, it was not accepted in 

that work, and must therefore be considered invalid according to Art. 34.1. The second appearance was in 

Stirp. agri. femsion. page 37 (June 1825) in the form of the combinations 'Micarea fuliginea and 'M. nigra'. 

Neither of these was accompanied by a description and the generic name was again invalidly published 

(Arts 32, 34.1(e)). 

When validly published, Micarea was introduced with four species: M. prasina, M. coccinea, M. 

fuliginea, and M. nigra. Korber (1855) emended the genus to include only M. prasina and is thereby 

considered to have lectotypified the genus on that species. See 'Excluded taxa' for notes on the remaining 

three original names. 

Thallus crustose or immersed in the substratum, effuse and often wide-spreading, never 

bordered by delimiting hypothalline lines. Superficial thallus consisting of ± spherical granules 

(goniocysts) up to c. 60 ±m diam; or convex to subglobose areolae which in some species may 

dissolve into soredia: or more rarely present as a thin ± smooth to rmose, or scurfy-granular 

crust. Thallus in section ecorticate (or weakly corticate in a few species), but sometimes (when 

areolate) covered by a hyaline amorphous layer, and outermost hyphae sometimes pigmented. 

Phycobiont 'grass-green'; cells usually thin-walled and c. 4-7 pm ('micareoid'), or more rarely 

larger ('non-micareoid' phycobiont types); a few species also have cephalodia containing Nostoc 

ox Stigonema. 

Apothecia small (mostly < 1 mm diam) , whitish or variously coloured (mostly grey , dull brown 

or blackish), epruinose, usually immarginate or ± so, adnate, sessile or rarely stipitate, convex 

to ± globose and often becoming tuberculate. Hymenium with amyloid gel-matrix. Asci clavate 

or cylindrical-clavate, of Lecanora-type, 8-spored. Spores hyaline, smooth-walled, variously 

shaped (ellipsoid, ovoid, fusiform, or acicular), usually less than 6 pm wide, simple to 

multiseptate but never muriform. Paraphyses few to numerous, septate, mostly branched 

(especially above), often anastomosing, mostly in range of 0-7-T7 pm wide at mid-hymenium;


apices never regularly clavate or capitate and never with a dark brown apical 'cap', but 

sometimes irregularly incrassate and sometimes with thickened pigmented walls in about upper 

5-15 ixm; in addition, some species have a few stout paraphyses (which are c. 2-3 /xm wide, and, 

in some cases, pigmented throughout their length) occurring as scattered individuals or small 

fascicles. Hypothecium (including subhymenium) variously pigmented, of interwoven hyphae 

that become outwardly orientated towards the hymenium, mixed with wider, short-celled 

ascogenous hyphae. Excipulum often absent or indistinct, if discernible then non-amyloid (or ± 

so) and composed of radiating branched and anastomosing paraphysis-like hyphae that become 

distinct and ± separate in K. 

Pycnidia often present, very varied in form from immersed to sessile or stalked, the stalks 

(pycnidiophores) sometimes branched. Pycnidial walls hyaline or pigmented. Conidiogenous 

cells ampulliform to cylindrical (sometimes with swollen base), phialidic, sometimes with 1-3 

proliferations. Conidia hyaline, smooth-walled, of three basic types: (i) microconidia - ± 

cylindrical, aseptate, eguttulate, in range 3-5-9x0-5-l fxm, borne in immersed to sessile 

pycnidia usually


species in at least one case. Micarea intrusa {q.v.) possibly provides a link between the two 

genera, and its inclusion in Micarea, rather than Scoliciosporum, is mainly on the grounds of 

spore-type. The differences between the two genera are admittedly unclear and ill-defined at the 

present time, and will certainly require re-appraisal as our knowledge of morphological, 

ontogenetic, and phylogenetic aspects in the Lecideaceae increases. 

A genus which may be closely related to both Micarea and Scoliciosporum is Strangospora 

Massal.; it shares with them unsophisticated apothecial and thallus structures, and similar 

ecological requirements. However, Strangospora differs in having polysporous asci with aseptate, 

globose spores, although the asci do appear to be of the Lecanora-typQ. The monotypic 

genus Steinia Korber is represented by S. geophana (Nyl.) Stein, whose small, black, immargin- 

ate apothecia occurring on shaded substrate (soil, rotten wood, rocks, stones, etc.) are easily 

confused in the field with those of some Micarea species. However, it is easily distinguished in 

microscopical preparations by its 16-spored asci with globose spores, and simple, slender 

paraphyses. In addition, its asci are not of the Lecanora-typQ, although their apices are each 

provided with a broad, amyloid plug. 

Micarea appears to have some affinities to Psilolechia Massal., and the two genera are 

compared in the discussion of Psilolechia clavulifera (p. 376). Micarea may also be close to the 

predominantly foliicolous Byssoloma Trevisan, but species of this latter genus have an excipulum 

of loosely woven, hyaline, pachydermatous hyphae which spread laterally to form a 

conspicuous white border to the apothecium. However, some specimens of B. subdiscordans 

(Nyl.) P. James, when growing on bark, mosses, or rocks, do not have conspicuous whitebordered 

apothecia, although the pachydermatous hyphae are clearly seen in sections of the 

apothecia. 

Lignicolous members of the 'Lecanora' symmicta group are occasionally confused with 

Micarea species on account of their general appearance and excipulum structure. Their 

excipular hyphae are radiating, richly branched and anastomosed and lax in K, although they 

contrast markedly with the simple or sparingly branched paraphyses and do not give the 

impression of being 'paraphysis-like'. This group further differs from Micarea in having dense 

epithecial granules which dissolve in K and a generally different suite of Uchen substances (which 

includes usnic acid and zeorin). Another seemingly distinctive group of species (but not yet 

afforded generic status) that bears some close resemblances to Micarea is that containing such 

species as Catillaria contristans, Lecidea limosa, and L. stenotera; for further discussion see 

account oi Micarea assimilata (p. 115). 

The apothecia of Vezdaea species are superficially similar to those of some species oi Micarea, 

but differ in that their tissues are not bound by a gelatinous matrix and their asci have uniformly 

amyloid walls and do not belong to the Lecanora-type; see Poelt & Dobbeler (1975). 

There has been some confusion between Micarea and genera such as Arthonia Ach., and 

Chrysothrix Mont., and also the Trapelia-\ike 'Lecidea' granulosa group. These all differ from 

Micarea in not having Lecanora-typc asci (see 'Excluded taxa'). 

Suprageneric considerations 

The formulation of a ± stable hierarchical classification of fungal (including lichen) taxa above 

the rank of genus is unUkely to be achieved for some decades. A suprageneric classification for 

the lichenized fungi has been attempted in recent years by, for example , Henssen & Jahns (1973) 

and Poelt (19746). These schemes are extremely useful bases for further study but, as admitted 

by their authors, contain very many uncertainties and points of conjecture, and should be 

regarded as being of a very provisional nature. 

Micarea has Lecanora-type asci and clearly belongs in the order Lecanorales and suborder 

Lecanorineae as defined by Poelt (19746). My belief is that Micarea should be placed in the 

Lecideaceae Chev. s. sir. (i.e. confined to species with Lecanora-type asci), with the provision 

that the real differences (if any) with that family and the Lecanoraceae Fee need to be explored. 

For the time being (at least) I cannot accept the 'Micareaceae Vezda ad int.' (Poelt, 19746: 627; 

Eriksson, 1981), although it may be possible to formally define this family name in the future as 

knowledge and taxonomic concepts in the Lecanorales increases and advances.

Infrageneric considerations 


I have given much thought to the possibihty of subdividing the genus Micarea into subgenera or 

sections. The genus includes several small groups of closely related species, but the affinities of 

many individual species are difficult to ascertain. I believe that a formal infrageneric classifica- 

tion (above the rank of species) based on current information would be unhkely to withstand the 

tests of time and serve httle useful purpose. However, that is not to say that such a classification 

will never be possible. The present study is mainly confined to the 45 European species of 

Micarea, but the genus is well represented in other parts of the world by some of the same, plus 

numerous additional species. With such considerations in mind I estimate that the genus, as 

currently circumscribed, contains about 100 species world-wide. A better understanding of 

extra-European species may increase the feasibility of subdividing Micarea. 

Some examples of small groups of apparently closely related species are given below. Within 

each group all the species have basically similar apothecial construction. For example. Group C 

all have medium to large sized apothecia, a ± well-developed excipulum, and numerous 

paraphyses which are of medium thickness and are richly branched (especially in the upper 

hymenium). Some supplementary common features and other notes are provided for each 

group - but for more detailed discussions see the taxonomic accounts of the relevant species. 

Phycobiont is 'micareoid' unless otherwise stated. See under 'Chemistry' for explanation of 

pigments. Abbreviations: AL, hyaUne amorphous covering layer; Exc, excipulum; Hym., 

hymenium; Hyp., hypothecium; Th., thallus; Pyc, pycnidia. 

A. M. lignaria - M. ternaria 

Th. areolate-type with AL. Th., Hym. and Pyc. with pigment A; Hyp. with dilute pigment A plus 

dilute dull brown pigment. Spores ± fusiform, 3- or more septate. Mesoconidial states very similar; 

M. lignaria also has micro- and macroconidial states. Chemistry: argopsin or xanthones (M. 

lignaria) or no substances {M. ternaria). 

B. M. leprosula-M. subleprosula 

Th. sorediate-type. Th. and Hym. with pigment A; Hyp. often with dilute dull brown pigment. 

Spores ± fusiform, 3- or more septate. Anamorphs unknown. Chemistry: argopsin -I- gyrophoric 

acid (M. leprosula) or alectorialic acid -I- accessory substances (M. subleprosula). Many affinities 

with Groups A and C. 

C. M. peliocarpa - M. alabastrites - M. cinerea 

Th. areolate-type with AL. Hym. , Th. and Pyc. of M. peliocarpa and M. cinerea with pigment A; 

all parts of M. alabastrites devoid of pigment. Spores ± fusiform, 3- or more septate. All with 

microconidia and curved or filiform macroconidia; mesoconidia unknown. Chemistry: all with 

gyrophoric acid. With Atlantic or sub-Atlantic distributions. Some affinities with Groups A, B, and 

D. 

D. M. denigrata - M. nitschkeana - M. globulosella 

Th. areolate-type but with no AL. Hym., Th. and Pyc. with pigment D; Hyp. hyaline. All with 

similar micro- and mesoconidial states; M. denigrata and M. nitschkeana also with ± identical 

macroconidial states. Chemistry: all with gyrophoric acid. Mainly corticolous or lignicolous. These 

three species appear to show a good example of an evolutionary progression: they are morphologi- 

cally and chemically ± identical except for the length and septation of spores. 

E. M. pycnidiophora - M. stipitata 

Th. areolate-type, but with no AL. Without pigments. Spores ± acicular, 3-7-septate. Pyc. 

stalked, with mesoconidia; micro- and macroconidia unknown. Chemistry: gyrophoric acid (Af. 

pycnidiophora) or no substances (M. stipitata). Corticolous and with eu-Atlantic distributions. 

F. M. elachista - M. rhabdogena 

Th. areolate-type with cortex and AL (M. elachista) or endoxylic (M. rhabdogena). Upper Hym. 

with pigment E; Pyc. with pigment D; Hyp. hyaline. Macroconidia unknown. Chemistry: no 

substances. Mainly lignicolous with boreal or boreal-continental distribution. 

G. M. botryoides-M. melaeniza 

Th. weakly areolate (with no AL) or with goniocysts (Af. botryoides) , or endoxylic {M. melaeniza). 

Hym. and Pyc. walls with pigment A; Hyp. and Pyc. stalks with pigment F. Paraphyses dimorphic. 

Pyc. stalked with mesoconidia; micro- and macroconidia unknown. Chemistry: no substances. This


group could possibly be extended to include M. anterior, and there are probably some affinities to 

species of Groups H and I. 

H. M. contexta - M. eximia - M. nigella - M. olivacea 

Th. endoxylic, or forming a thin crust, or weakly areolate. Pigments A and (or) B in various 

locations. Spores simple or 1-septate. Paraphyses dimorphic. Pyc. black, immersed to sessile, or 

stalked (A/, nigella). Macroconidia unknown. Chemistry: no substances. Mainly lignicolous and 

confined (?) to north-west Europe. Probably close to Group G. M. melaena may belong here but 

has: rather large apothecia, pigment C (usually present in hypothecium), an often well-developed, 

areolate thallus containing gyrophoric acid, and an ability to produce macroconidia; it may provide 

a link between Group H and Groups A-D. 

I. M. bauschiana - M. sylvicola - M. tuberculata - M. lutulata 

Th. weakly areolate, ± smooth or scurfy. Phycobiont non-micareoid. Pigmentation variable, 

involving pigments A, B, C, and F; pigment D never present. Spores small, simple, or p.p. 

1-septate (M. tuberculata). Paraphyses dimorphic. Pyc. ± immersed. Macroconidia unknown. 

Chemistry: no substances. Mainly found in dry underhangs in the Micareetum sylvicolae. This 

group is almost worthy of subgeneric status but it shows some affinities to Group G, and other 

species, such as M. lithinella. 

J. M. assimilata - M. incrassata - M. subviolascens - M. melaenida - M. crassipes 

Th. areolate-type, sometimes with AL. Pigmentation variable, involving pigments A, B, C, D, and 

E; Hyp. always dark-coloured. Spores rather large, mostly ellipsoid or oblong-elhpsoid and simple 

or 1-septate. Paraphyses rather stout and sparingly branched. Cephalodia found in first three 

species. Pyc. immersed. Macroconidia unknown. Chemistry: no substances. Mainly terricolous or 

muscicolous (never corticolous or lignicolous), and with a ± arctic or arctic-alpine distribution 

(except M. melaenida). M. crassipes is rather anomalous here because of its very well-developed 

excipulum and turbinate or stipitate apothecia. 

K. M. prasina - M. hedlundii - 'Lecidea' levicula [from Cuba] 

Th. goniocyst-type. Hym., Pyc. and Th. often with pigment D; Hyp. hyaUne; one species (M 

hedlundii) with pigment H. Spores mostly simple or 1-septate. Pyc. immersed to sessile, or stalked 

and tomentose {M. hedlundii). Micro- and (or) mesoconidia produced; macroconidia unknown. 

Chemistry: variable, including 'prasina unknowns' and gyrophoric acid. M. misella and M. 

melanobola have some affinities to this group, but also with Group D. M. synotheoides may belong 

near here. 

Keys to species 

Guide to keys and identifications 

Two keys are provided, the first to specimens with apothecia, the second to specimens without 

apothecia (although often with pycnidia). The keys are for European species, but all those 

known to me from cool-temperate, boreal, and arctic regions of North America are, by chance, 

included. 

The nature of pigmentation and colour reactions observed in anatomical sections is important 

for the identification of Micarea species. Many of the subtle yet significant colour hues and 

reactions are obscured by the yellowish illumination given by most bulbs, and it is recommended 

that a microscope be fitted with a 'daylight' bulb or a blue filter. The preceding sections on 

'Morphology' and 'Chemistry' should be read before attempting to use the keys. 

Particular care should be taken in determining C reactions with apothecial sections. The straw 

to dull olivaceous, K+ violet pigment found, for example, in M. prasina, M. synotheoides, M. 

denigrata, and M. nitschkeana, also reacts C+ violet (persistent); this reaction tends to 

mask the C+ orange-red (quickly fading) reaction due to gyrophoric acid and usually obtainable 

in the last two, aforementioned species. However, the C+ violet reaction is mostly confined to 

the upper part of the hymenium, whereas the C+ orange-red reaction occurs in all parts of the 

apothecium. To carry out the C test cut a hand section of an apothecium and mount in a drop of 

water near the edge of the cover-slip; lay a piece of tissue-paper over the cover slip to take up any 

excess water; place the slide preparation on the microscope stage and focus at about x 100 to 

x200 and note any pigmentation; then, apply a drop of C by the edge of the cover-slip nearest


the section. A positive reaction due to gyrophoric acid is seen as an orange-red front moving 

across the section, with the colour quickly fading in its wake. If the concentration of gyrophoric 

acid is very high than the reaction may be more immediate and intense, but, nevertheless, the 

colour quickly fades away within a few seconds. A similar procedure should be followed when 

testing for other colour changes. 

The identification of Micarea species usually requires accurate measurement of the dimen- 

sions of spores, conidia, and paraphyses, such that a microscope with a good resolution at 

X 1000, coupled with a carefully calibrated measuring eyepiece, is invaluable, and in many cases 

essential. 

Spores should be observed and measured in 10% KOH; in most cases this solution will render 

any septa visible. If, however, the spore contents are difficult to clear, septa can be discerned by 

preparing an apothecial section or squash in LCB and heating (just to boiHng) over a spirit lamp; 

any septa should then be clearly seen, especially under oil-immersion (xlOOO). 

Conidia should always be examined under oil-immersion (at least xlOOO). Preparations can 

be made in 10% KOH, but better resolution is obtained using ammoniacal erythrosin. In water, 

small conidia are prone to 'Brownian movement', making alignment with the measuring 

graticule a frustrating task! If conidia and conidiogenous cells are to be examined in a squash 

preparation it is often helpful to soak an intact pycnidium in a small drop of 10% KOH on a shde 

for about a minute; excess KOH is then soaked up with the edge of a piece of tissue-paper or 

filter-paper and the squash prepared under a cover-slip in ammoniacal erythrosin. The KOH 

softens the pycnidial wall and does not alter the effectiveness of the erythrosin. 

Fine measurements, such as the width of spores, conidia, hyphae, and paraphyses should not 

be made in LCB, this solution often causing much shrinkage; with hyphae and paraphyses the 

cytoplasm (lumina) is intensely stained, but the delimitation of their outer walls is often difficult 

to ascertain. 

Key to European species 

la Hymenium, atleast in upper part, dull greenish or brownish in water, K-l- violet [pigment D].. 2 

lb Hymenium variously coloured or hyaline, not turning to violet in K (if purple in K, then 

pigment already purplish in water [pigment C] 12 

2a (la) Hypothecium dark purple-brown, K+ purple intensifying or K+ green in upper part. 

Saxicolous 40. M. subviolascens (p. 185) 

2b Hypothecium hyaline or pale . Usually corticolous or lignicolous 3 

3a (2b) Spores mostly 3 (or more)-septate , or over 15 /u,m long 4 

3b Spores mostly simple or 1-septate (2- or 3-septate spores if present very rare) , mostly less than 

15 /am long 

4a (3a) Apothecia sections and/or thallus C + orange-red (gyrophoric acid) 5 

4b Apothecia sections and thallus C- [note: the K+ violet pigment also reacts C+ violet], 

gyrophoric acid absent 

5a (4a) Spores 0-l(-2)-septate, few exceeding 16 /xm in length. Usually lignicolous. Rare forms 

of this very variable species 11. M. denigrata (p. 127) 

5b Spores mostly 3 (or more)-septate, or many exceeding 16 /am in length. Usually on bark or 

twigs 

6 

6a (5b) Spores (l-)3-septate, fusiform, often curved, 10-17(-19)x2-5-3(-3-5) /am. Microconidia 

(4-7-)5-5-7-5x0-8 /am; mesoconidia (3-)3-5-5(-5-7)x 1-1-5 /am; macroconidia some- 

times present, curved 30. M. nitschkeana (p. 165) 

6b Spores 0-3(-6)-septate, acicular, 13-26xl-5-2-5(-3) /am. Microconidia 3-8-5 xO-8-1 /am; 

mesoconidia 3-6-5-3 x 1-1-4 /am; macroconidia unknown 14. M. globulosella (p. 134) 

8 

7


7a (4b) Thallus dull grey-green (never whitish) to dark olivaceous or blackish, of discrete or 

coalescing granular-areolae c. 20-70 /xm diam, ± gelatinous when wet. Phycobiont 

micareoid, cells 4-8 /im diam. Apothecia grey- or brown-black, epruinose. Spores 

l-5(-ll)-septate, 14-35(^3) xl-8-2-5(-3) fim. Microconidia and/or mesoconidia often 

present, both exceeding 3-7 fxtn in length. An oceanic species on acid bark (sometimes 

over bryophytes) 

42. M. synotheoides (p. 188) 

7b Thallus whitish, tinged grey or green, ± endophloeodal or thin and smooth to ± verrucose, 

often rimose; never gelatinous. Phycobiont not micareoid, cells 8-16 ixm diam. Apothecia 

grey to black, often with whitish pruina. Spores (l-)3-7-septate, 17-26x1 •5-2-5 )u,m, 

Excipulum hyphae coherent in K. Conidia of one type only, 2-8-3-5xl-l-4 /xm. Widespread 

species on rather basic bark , rarely on lignum Bacidia beckhausii 

8a (3b, 31a) Thallus of pale to dark green goniocysts c. 12-40(-60) fxm diam, often ± gelatinous 

when wet, C-, gyrophoric acid absent or detectable in trace amounts (?contaminant) by 

t.l.c 

8b Thallus endoxylic or of whitish or grey granular-areolae c. 60-200 fim diam, not appearing 

gelatinous when wet, CorC-l- orange-red , gyrophoric acid often present 11 

9a (8a) Goniocysts K- or hyphae violaceous in K, pigment never oily. Pycnidia immersed or 

sessile, not tomentose 10 

9b Goniocysts containing purple oily substance in K. Pycnidia numerous, distinctly stalked, 

brown with white tomentum; containing mesoconidia (4-)4-5-5-5(-6)xl-3-l-7 /xm. 

Apothecia usually few or absent, brown, soon tuberculate. On soft lignum of conifers, 

rare 

15. M. hedlundii (p. 135) 

10a (9b) Apothecia dark grey to black, small, 0-1-0-24 mm diam. Paraphyses with dark apical 

walls giving a distinct dark green, K+ violet epithecium. Spores 0-1-septate, 7-9-7x 

2-5-3-3 /xm. Microconidia 4-5-5 xO-7-0-8 /xm; mesoconidia 3-3-4-5x1-1 -3 /xm. 

10b 

Thallus indistinct, of scattered olivaceous goniocysts. Rare, (?) confined to Picea bark in S. 

Finland 

25. M. melanobola (p. 156) 

Apothecia white to grey, sometimes blackish, 0-1-0-4 mm diam. Without sharply delimited 

epithecium, the olivaceous, K-l- violet pigment mostly confined to the gel-matrix and 

± diffuse although often more concentrated towards upper part of hymenium. 

Spores variable, 0-l(-3)-septate, 8-14(-17)x2-3-4(-5) /xm. Microconidia (5-)5-5-8x 

0-7-1 /xm; mesoconidia 3-5-4-7x1-4-1-8 /xm. Thallus of thinly 

goniocysts, sometimes forming a thick crust. Common and 

scattered to coherent 

widespread on bark, 

lignum and turf (especially coastal), rare on rock; morphologically and chemically very 

variable 

34. M. prasina (p. 173) 

11a (8b) Apothecia grey to black, sometimes pallid or brown (shade forms), 0-1-0-5 mm diam. 

Thallus of distinct whitish areolae, or dark and scurfy (parasitized), occasionally endoxylic. 

Spores mostly 1-septate, often ± curved, (7-)9-16(-18)x2-3-3(-3-5) /xm. Paraphyses 

numerous, c. 1-1-5 /xm wide. Pycnidia immersed or emergent, never distinctly stalked. 

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- 

dia curved, 12-24x1 /xm. Apothecia and thallus sections usually C-l- orange-red (gyrophoric 

acid) 

ll.M. denigrata(p. 127) 

lib Apothecia black (very rarely paler), smaller, 0-1-0-3 mm diam. Thallus usually endoxylic, 

rarely forming whitish areolae. Spores mostly simple, 1-septate spores rarely numerous, 

never curved, 6-5-9-5x2-3(3-7) /xm. Paraphyses scanty, thin, 0-5-0-8 /xm wide. EndoxyUc 

forms usually with black, stalked pycnidia containing mesoconidia, 3-5-5x1-1-5 /xm; 

microconidia 3-8-6x0-7-0-8 /xm, borne in immersed pycnidia; macroconidia unknown. 

Apothecia sections usually C-, very rarely C-l- orange-red; thallus when superficial and 

well developed C+ orange-red (gyrophoric acid) 26. M. misella (p. 158) 

12a (lb) Spores acicular or sigmoid-curved (mostly over 18 /xm long and under 3-5 /xm wide at 

maturity) 

13 

12b Spores not acicular or sigmoid-curved 15 

9


13a (12a) Apothecia whitish, without pigmentation, epruinose, C+ red or C-. Phycobiont 

micareoid , cells 4-7 /xm diam . Pycnidia always abundant, sessile or stalked, whitish 14 

13b Apothecia dark coloured (with brownish or greenish pigment in upper hymenium); if 

whitish then pruinose (epithecium finely granular) or not. Phycobiont not micareoid, 

many cells over 8 ^tm diam. Pycnidia (?) unknown; if present, then never conspicuous or 

stalked Scoliciosporum species 

[see Poelt & Vezda (1981)] 

14a (13a) Apothecia and pycnidia C+ red (gyrophoric acid). Pycnidial structures mostly 

unbranched, 0- 1-0-3 mm tall; containing mesoconidia 4-6x1-1-5 fim 

[if spores fusiform and pycnidia immersed, see 23] 

35. M. pycnidiophora (p. 179) 

14b Apothecia and pycnidia C- . Pycnidial structures often branched, 0-3-0-8 mm tall; containing 

mesoconidia 6-8x1-1-8 )u,m 37. M. stipitata (p. 182) 

15a (12b) Mature spores 3- or more septate, mostly over 15 fim long 16 

15b Mature spores simple or 1-septate 28 

16a (15a) Upper hymenium with distinct epithecial layer of dense fuscous-brown pigment (without 

green or purple tinge), K+ dissolving into a brown solution. Spors 0-l(— 3)-septate, 

(9-)ll-15(-20)x2-3-5/Am. Hypotheciumpale. Usually lignicolous 12. M. elachista (p. 131) 

16b Upper hymenium otherwise, if brownish then pigment not dissolving into a brown solution. 

Hypothecium hyaline , pale or dark. Substrata various 17 

17a (16b) Upper hymenium purple, K-l- intensifying. Hypothecium dark purple-brown. Rare 

form of 22. M. melaena (p. 150) 

17b Upper hymenium hyaline , greenish (sometimes tinged brown) or aeruginose 18 

18a (17b) Hypothecium dark in thin section 19 

18b Hypothecium hyaline or with dilute greenish or brownish tinge 20 

19a (18a) Hypothecium mottled reddish brown, K-, HNO3- or + orange-brown. Upper 

hymenium greenish, sometimes brownish in part. Hyaline paraphyses 1-1-5 /nm wide. 

Thallus always C- (gyrophoric acid absent) ;terricolous 45. M. turfosa (p. 194) 

19b Hypothecium dark purplish brown, K-l- green or K-l- purple intensifying (often a mixture of 

both reactions, HNO3-I- purple-red. Hyaline paraphyses 0-8-1 /urn wide. Thallus when well 

developed C-f- red (gyrophoric acid) 22. M. melaena (p. 150) 

20a (18b) Apothecia whitish, without pigmentation, C-l- red (gyrophoric acid). Spores mostly 

20b 

3-septate 

Apothecia pale grey to black, often with a greenish tinge; if whitish then spores mostly 

7-septate (M. cinerea). Upper hymenium greenish to aeruginose. Apothecia sections C-, 

or C-l- orange-red (gyrophoric acid). Spores 3- or more septate 

21a (20a) Apothecia usually numerous and crowded. Spores 3(-5)-septate, (16-)17-26(-28)x 

4-5(-6) ^tm. On bark and lignum, often over bryophytes, in oceanic areas 

2. M. alabastrites (p. 110) 

21b Apothecia usually dispersed and usually with some hint of pigmentation. Spores 

(l-)3(-5)-septate, (ll-)15-23(-24)x3-5(-6) /am. Shade forms, occurring on bark of 

old trees (especially Quercus) or on rocks , of this widespread and variable species of various 

substrata (see 27a) 

21 

22 

33. M. peliocarpa (p. 169) 

22a (20b) Thallus of fragile (easily broken with point of dissecting needle) , ash-grey to grey-brown 

granular-areolae, often dissolving to form sorediate patches, PD-I- yellow or red. Usually 

sterile 

22b Thallus of firm areolae or granules, or endoxylic, PD- or PD4- red (M. lignaria). Apothecia 

usually numerous 

23a (22a) Thallus C-l- red, PD-I- red (gyrophoric acid and argopsin). Spores (l-)3-septate, 

14-26(-29)x4-5-5/Ltm 

18. M. leprosula (p. 140) 

23b Thallus Cf+ red, KC-I- red, PD+ deep yellow (alectorialic acid). Spores 3-7(-9)-septate, 

35-45(-60) X 5-6-5 /im 

38. M. subleprosula (p. 182) 

23 

24


24a (22b) Thallus PD+ red and C- (argopsin), or PD- and C+ persistent orange (xanthone); 

gyrophoric acid absent. Apothecia black, ± globose, sessile or occasionally stipitate. Upper 

hymenium green (sometimes brownish in reflexed parts). Hypothecium pale, dilute 

greenish or brownish in upper part . Spores 3-7-septate , 16-36(-38) x 4-6(-7) fixn 25 

24b Thallus PD— ; thallus and/or apothecia sections C+ red (soon fading; gyrophoric acid) or C— 

(no substances) . Apothecia often adnate at first 26 

25a (24a) Areolae C-, PD+ red (argopsin), usually whitish or pale grey. Widespread and 

common, especially in upland areas 19a. M. lignaria var. Iignaria(p. 142) 

25b Areolae C+ persistent orange, PD— (xanthone), usually whitish with yellowish (isabeUine) 

tinge. Local, confined to high rainfall areas, often with var. lignaria 

19b. M. lignaria var. endoleuca (p. 146) 

26a (24b) Spores (3-)5-7-septate, (19-)23-34(-34(-38)x4-5-6 fxm. Macroconidia filiform, flexuose, 

50-1 10x1 jxm. Thallus and apothecia sections C+ red (gyrophoric acid) 

7. M. cinerea (p. 121) 

26b Spores mostly 3-septate (5-septate spores absent or in very low frequency), less than 25 (xm 

long. Macroconidia less than 50 /i-m long. Thallus and apothecia sections C+ red or C- 27 

27a (26b) Thallus and apothecia sections C + red (gyrophoric acid) . Hypothecium usually hyaline 

Microconidia (5-)6-7(-7-7)x0-4-0-7 /xm; mesoconidia unknown; macroconidia curved, 

27b 

21^0(^7) X 1-1 -5 fxm. Widespread on various substrata 

Thallus and apothecia sections C— (no substances with t.l.c). 

33. M. peliocarpa (p. 169) 

Hypothecium with greenbrown 

tinge (as in M. lignaria). Microconidia and macroconidia unknown; mesoconidia 

4-6-6-3-1 -2-1 -7 fxm. On turf in the Arctic; possibly occurring in western Britain on rocks in 

coastal districts as a form with a ± obsolete thallus 43. M. ternaria (p. 190) 

28a (15b) Thallus of well developed, whitish to grey-brown, convex granular-areolae, 0-08- 

0-4 mm diam, with intermixed brown cephalodia containing Nostoc or Stigonema. Apothecia 

black, convex to subglobose, never markedly constricted below. Spores simple, or 

l(-2)-septate spores intermixed, 10-17(-19)x3-5 fxm. Upper hymenium green, K-. 

Hypothecium dark. Over bryophytes, plant debris or light soil amongst rocks or on exposed 

turf in montane or Arctic regions 29 

28b Without the above combination of characters ; never with cephalodia 30 

29a (28a) Hypothecium reddish brown, K— , HNO3-I- bright orange-brown (no purpUsh tinge). 

Thallus usually grey-brown. Spores simple or often 1-septate (rarely 2-septate) 

16. M. incrassata (p. 137) 

29b Hypothecium purple-brown, K-l- purple intensifying, HNO3-I- purple-red. Thallus usually 

whitish or brownish white . Spores mostly simple , septate spores very rare 

[if young apothecia turbinate and thallus finely granular, see 51b] 

4. M. assimilata (p. 114) 

30a (28b) Apothecia whitish, pallid or dull reddish, without distinct pigmentation in section. 

Spores mostly 1-septate and over 9 /xm long. Phycobiont micareoid. On bark or lignum or 

over bryophytes thereon 31 

30b Apothecia coloured with obvious pigmentation in section; if not then, spores simple and/or 

smaller, or on rock , or phycobiont not micareoid 33 

31a (30a) Thallus comprised of goniocysts/or apothecia sections C-l- red (gyrophoric acid) and 

spores less than 3 fim wide . Shade 

forms 8 

31b Thallus not comprised of goniocysts, endoxylic or weakly areolate, or scurfy granular, or 

rimose, sometimes with waxy appearance. All parts C— (gyrophoric acid absent). Spores 

often over 3 fim wide 32 

.


32a (31b) Apothecia pale reddish or orange-brown, small, 0-15-0-35 mm diam, sometimes 

tuberculate and up to 0-6 mm diam, but never forming large nodulose clusters. 

Spores ovoid-fusiform, often slightly curved, (0-)l(-3)-septate, 9-16x2-5-4-5 /xm. Pale 

reddish brown, stalked pycnidia often present. Thallus usually poorly developed, never 

waxy 3. M. anterior (p. 112) 

32b Apothecia pallid, at first ± plane and adnate, 0-2-0-4(-0-6) mm diam, later often coalescing to 

form large nodulose clusters up to 2 mm wide. Spores ellipsoid, ovoid or oblong, never 

curved, (0-)l -septate, 9-16x3-5 ixm. Without stalked pycnidia, but white, convex sporodochia 

(0-1-0-25 mm diam) usually present, bearing simple, ellipsoid macroconidia, 

6-10x2-3 /um l.M. adnata (p. 108) 

33a (30b) Thallus and apothecia sections C-l- red (? gyrophoric acid). Epithecium fuscous-brown. 

Hypothecium pale. Spores markedly curved, 1-septate , 9-12 x 2-5^ fim . On rock 

10. M. curvata (p. 126) 

33b Thallus and apothecia sections C—; without remaining combination of characters 34 

34a (33b) Upper hymenium with well defined fuscous-brown epithecium, pigment unchanged in 

colour but dissolving into solutions in K. Hypothecium hyaline to dilute yellowish brown. 

Phycobiont micareoid. Mostly on lignum or old bark 35 

34b Upper hymenium otherwise; if fuscous-brown then pigment not dissolving in K and plant not 

on bark or lignum. Hypothecium hyaline, pale or dark. Phycobiont micareoid or not. 

Substrata various 36 

35a (34a) Spores 0-l(-3)-septate, (9-)ll-15(-19)x 2-3-5 /am. Thallus of dispersed to contiguous, 

whitish, grey-brown or olivaceous-brown, convex to ± globose areolae. On lignum or old 

bark, very rarely on rock 12. M. elachista . (p 131 

35b Spores 0(-l)-septate, 6-9-5 x 1 -5-2-3 /xm. Thallus endoxyUc. On lignum 

36. M. rhabdogena (p. 181) 

36a (34b) Hypothecium hyaHne, yellowish, dilute orange-brown or dilute greenish, not blackish 

in thick sections 37 

36b Hypothecium dark throughout, often blackish in thick section 42 

37a (36a) Apothecia dark brown or black. Spores 0-1-septate, c. 9-14x4—6 ± m. On exposed 

rocks 38 

37b Apothecia pale; if dark then spores simple and/or smaller. In sheltered situations on shaded 

rocks, exposed tree-roots etc 39 

38a (37a) Upper hymenium brown. Phycobiont micareoid. Pycnidia usually present, with either 

helicoid macroconidia or cylindrical microconidia 39. M. subnigrata (p. 183) 

38b Upper hymenium green. Phycobiont not micareoid, thick-walled, 7-21 /xm diam. Pycnidia 

unknown 17. M. intrusa (p. 138) 

39a (37b) Spores small, in range, 4-8-5x1-2-5 /xm, single or 1-septate. Apothecia small, mostly 

less than 0-3 mm diam 40 

39b Spores larger, c. 7-10x3^-5 fim, simple. Apothecia usually larger, often more than 0-3 mm 

diam 41 

40a (39a) Spores mostly 1-septate, 6-8-5x1-5-2-5 /xm. Apothecia orange-brown or reddish 

brown . Hypothecium 

dilute orange-brown . Phycobiont (?) micareoid , cells 4-7 /am 

28. M. myriocarpa (p. 161) 

40b Spores simple, 3-6x1-2 /am. Apothecia blue-grey to blackish. Hypothecium dilute green. 

Phycobiont cells 5-12(-18)x 3-8 /am, often in pairs or short chains Psilolechia clavuiifera (p. 201) 

41a (39b) Phycobiont not micareoid, cells 5-12 /am diam. Apothecia pallid to black. Upper 

hymenium hyaline to dark green or aeruginose. Hypothecium hyaline, or dilute greenish in 

upper parts 5. M. bauschiana (p. 117) 

41b Phycobiont micareoid, cells 4-8 /am diam. Apothecia pallid, dull yellowish-orange to reddish 

brown. Upper hymenium hyaline. Hypothecium straw-yellow to dilute orange-brown 

20. M. lithinella (p. 147) 

42a (36b) Hypothecium fuscous or ± reddish brown, without distinct purple or greenish tinges in 

water or K, HNO3- or -I- bright orange-brown, never -I- purple-red 43 

42b Hypothecium with distinct purple or greenish tinge in water and/or K, HN03-F purple-red 50 

)


43a (42a) Pycnidia conspicuous, black , sessile or stalked 44 

43b Pycnidia inconspicuous and immersed, or absent 45 

44a (43a) Spores simple, 5-9x2-5-3-8 /xm. Conidia 2-6-3-6X 1-1 -3 fim. On lignum 

24. M. melaeniza (p. 155) 

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, 

decaying bryophytes, exposed tree roots or loose stones , rarely on bark or lignum 

6. M. botryoides (p. 118) 

45a (43b) On rocks, loose stones, consolidated soil, dry bryophyte mats or exposed tree roots in 

underhangs, or on lignum 46 

45b On mineral soil, or over decaying bryophytes or plant debris on the ground; never in 

underhangs 48 

46a (45a) Spores mostly 1-septate, 6-8-5x1-5-2-5 /xm. Hymenium hyaline or tinged orangebrown, 

never greenish. Hypothecium never blackish 28. M. myriocarpa (p. 161) 

46b Spores simple. Hymenium often greenish or ± aeruginose. Hypothecium blackish in thick 

section 47 

47a (46b) On sheltered rocks, rarely on exposed tree roots. Apothecia convex-globose, often 

tuberculate. Spores 6-8(-9)x2-3(-3-4) /xm. Hypothecium c. 120-360 fxm tall. Phycobiont 

not micareoid, cells 5-12 /xmdiam 21. M. lutulata (p. 148) 

47b On Hgnum, rarely on rock. Apothecia convex-adnate, never tuberculate. Spores 9-12x 

4-5 /im. Hypothecium c. 70-120 fim tall. Phycobiont micareoid, cells 4-7 fxm diam 

27. M. muhrii (p. 160) 

48a (45b) Spores simple , small , 6-9 - 5 x 3-4 /am . Upper hymenium brown without green or purple 

tinge in K. On exposed soil in woodland clearings (? sites of old bonfires) 

32. M. osloensis (p. 169) 

48b Spores larger, mostly over 10 ^tm long. Upper hymenium usually with greenish or purple tinge 

inK 49 

49a (48b) Thallus blackish. Upper hymenium usually with green tinge in K. Spores simple, or 

often becoming 3-septate (10-)12-21(-25)x(3-5-)4-5 /xm. On exposed soil or truf on 

mountain summits or exposed ridges, or at lower altitudes in boreal regions 45. M. turfosa (p. 194) 

49b Thallus whitish. Upper hymenium with purple tinge in K. Spores 0-1-septate, (7-)9-5-15x 

3-4(-4-5)/am. On fine-sandy or argillaceous soil 23. M. melaenida (p. 154) 

50a (42b) Thallus terricolous or muscicolous. Spores simple or 1-septate, mostly in range 

9-19x3-4-5 /am. Paraphyses numerous, simple or sparingly branched, 1-1-5 /am wide 51 

50b Thallus saxicolous, lignicolous or corticolous, not terricolous or if so (anomalous occurrences) 

then spores smaller or paraphyses thinner and much branched 52 

51a (50a) On fine sandy or argillaceous soils, never muscicolous or on plant debris; at low 

altitudes. Apothecia immarginate, convex to hemisphaerical, adnate when young; in 

section without greenish pigmentation in K. Spores 1-septate, (7-)9-5-15 x 3-4 /am 

23. M. melaenida (p. 154) 

51b On bryophytes or plant debris, sometimes spreading on to sandy soil; in Arctic or Alpine 

situations. Apothecia, at least when young, marginate, and markedly constricted below, 

turbinate or short-stalked; in section (especially excipulum) with green pigmentation 

(often mixed with purple pigmentation) in K. Spores mostly simple, (9-)10-19(-21)x 

(2-5-)3-4-5/am 9. M. crassipes (p. 125) 

52a (50b) Phycobiont not micareoid, cells 5-12 /am or more in diam. On shaded rocks in humid 

places (narrow ravines or woodlands), or in sheltered underhangs on rock, loose stones, 

exposed roots or consolidated soil ; rarely on lignum (old fence posts) 53 

52b Phycobiont micareoid , cells 4-8 /am diam . Usually on lignum , rarely on shaded rock 56 

53a (52a) Hypothecium dark reddish brown, K- or K± red intensifying, but not turning 

purplish or greenish. Hymenium hyaline, dilute green to aeruginose. Spores 6-8(-9)x 

2-3(-3-4)/am 21. M. lutulata (p. 148) 

53b Hypothecium distinctly greenish in K, more rarely purplish in part or in whole 54


54a (53b) Spores ellipsoid or ovoid-ellipsoid, (6-)7-10x(2-3-)3-4-5 fxm. Apothecia 0-2-0-5 mm 

diam, or reaching 1-2 mm when tuberculate. Conidia 3-8-6(-6-6) x 1-1 •7(-2) fjivn 

41.M. syIvicoIa(p. 186) 

54b Spores ovoid, oblong-ovoid or dacryoid, never ellipsoid, smaller, mostly less than 8 /u,m long 

and 2-4 fxm wide. Non-tuberculate apothecia usually smaller, rarely exceeding 0-3 mm 

diam. Conidia, if present, shorter 55 

55a (54b) Spores 0-1-septate, 5-8(-9) x 1 •5-2-5 fxm. Hypothecium much darker than hymenium. 

Pycnidia often present, 60-120 ^tm diam; wall dark green. Conidia 3-4-3x 1-1-4 fxm. 

Phycobiont cells ± globose, c. 5-12 /xm diam, never in short chains.. 44. M. tuberculata (p. 192) 

[if similar but phycobiont micareoid, see 58a] 

55b Spores simple, 3-6xl-l-7(-2) fim. Hypothecium concolorous with, or paler than, hymenium. 

Pycnidia unknown. Phycobiont cells ± globose, eUipsoid or oblong, 5-12(-18)x 

3-8 fim, often in pairs or short chains Psilolechia clavulifera (p. 201) 

56a (52b) Spores simple, 6-5-10x2-5-3-7 /xm. Sessile or stalked, black pycnidia present. 

Hypothecium purple-brown, K-l- green 29. M. nigella (p. 163) 

56b Spores (0-)l -septate. Pycnidia innate, never stalked 57 

57a (56b) Spores narrow, oblong-fusiform, 9-14(-16)x 1-8-2-5 /u.m. Hypothecium purple-brown, 

K-l- green. Hymenium bright green in upper part. HyaHne paraphyses rather scanty, thin, 

0-7-0-8 /Am wide. Pycnidia with mesoconidia3-9-5-5xl-l-4/Ltm 13. M. exima (p. 134) 

57b Spores broader , 2 - 5 fim or more in width , oblong with ± rounded apices , or ovoid 58 

58a (57b) Hyahne paraphyses numerous, 1-1-2 ^tm wide. Hypothecium dark sordid-olivaceous or 

ohve-brown, without any purple tinge in water or K. Hymenium oHvaceous green. Spores 

oblong or ovoid-oblong, (0-)l -septate, (7-)9-12-3x 2-5-3-5 /u,m. Pycnidia with mesoconidia3-4-2x 

1-1-3 /xm 31. M. olivacea (p. 167) 

58b Hyaline paraphyses scanty or numerous, thin, 0-6-1 /xm wide. Hypothecium often with purple 

tinge in water or K, green and purple pigments often intermixed 59 

59a (58b) Spores ovoid or oblong ovoid, 1-septate, 7-13(-14)x(2-3-)3^-5 fim. Apothecia 

0-1-0-2 /xm diam, or reaching 0-3 mm when tuberculate. Thallus endoxylic. Microconidia 

4-5x0-8-1 /am; mesoconidia3-8^-7x 1-3-1-8 />tm;macroconidia unknown 

8. M. contexta (p. 124) 

59b Spores oblong or ovoid oblong, at first 1-septate, becoming 3-septate at maturity. Apothecia 

0-12-0-4 mm diam, or reaching 0-5 mm when tuberculate. Thallus superficial, containing 

gyrophoric acid when well developed. Microconidia 4-8-7x0-8-1 /xm; mesoconidia unknown; 

macroconidia curved, 18-33 x 1-1 -5 /im. Immature forms 22. M. melaena (p. 150) 

Key to European species that may occur without apothecia 

Notes: all species included here have micareoid algae; all species with stalked pycnidia are 

included here. 

la Thallus of fragile ash-grey or grey-brown granular-areolae often dissolving or eroding to form 

sorediate patches; PD -I- yellow or red . Pycnidia absent 2 

lb Thallus otherwise , PD - . Pycnidia usually present and conspicuous ( x 20 lens) 3 

2a (la) Thallus C-l- red, PD -I- red (gyrophoric acid and argopsin) 18. M. leprosula (p. 140) 

2b Thallus C+ red (often faint), KC-I- red, PD+ deep yellow (alectorialic acid) 

38. M. subleprosula (p. 182) 

3a (lb) With stalked or sessile pycnidia containing mesoconidia (smaller ± immersed pycnidia 

containing microconidia sometimes present) ; never with curved or flexuose macroconidia 4 

3b Pycnidia innate sometimes becoming emergent with gaping ostioles, or with small sessile 

apothecia-Hke sporodochia; pycnidia sometimes containing curved or flexuose macroconi- 

dia 

4a (3a) Pycnidia black 

-^ 

4b Pycnidia whitish to reddish brown, never black ° 

5a (4a) Pycnidial wall oHvaceous brown, K-l- violet. Mesoconidia 3-5-5 x 1-1 -5(-l -7) fim 

5b Pycnidial wall K- or K-l- olivaceous 

11 

26. M. misella (p. 158) 

"


6a (5b) Pycnidial wall and stalk dark purple-brown, K-l- dark green. Mesoconidia 3-4-4-3X 

l-2-l-6)u,m 29. M. nigella (p. 163) 

6b Pycnidial wall dark olivaceous, K— ; stalk fuscous or reddish-brown, K— 7 

7a (6b) Mesoconidia 3-5-4-8X 1-1-5 /xm. On bryophytes, shaded rocks and stones or exposed 

roots, rarely on lignum 6. M. botryoides (p. 118) 

7b Mesoconidia2-6-3-6xl-l-3/Lim. Onlignum 24. M. melaeniza (p. 155) 

8a (4b) Thallus of pale to dark green goniocysts, 12-40 /x.m diam; goniocysts containing purple 

oily substance in K. Pycnidia brown with white tomentum. Mesoconidia (4-)4-5(-6)x 

1-3-1-7/xm 15. M. hedlundii (p. 135) 

8b Thallus indistinctly areolate or rimose, or endoxylic, without goniocysts, K— . Pycnidia 

without tomentum 9 

9a (8b) Pycnidia whitish, C-l- red (gyrophoric acid) . Mesoconidia 

4—6 x 1-1-5 /im 

35. M. pycnidiophora (p. 179) 

9b Pycnidia C— , whitish or reddish brown 10 

10a (9b) Pycnidia whitish, often branched. Mesoconidia 6-8x1-1-8 ^tm. Usually on bark, in 

oceanic areas 37. M. stipitata (p. 182) 

10b Pycnidia reddish brown, rarely branched. Mesoconidia 3-5-4-5 x 1-2-1-6 fim. On lignum in 

boreal regions 3. M. anterior (p. 112) 

11a (3b) With whitish, cushion-like, apothecia-like sporodochia c. 0-1-0-25 mm diam, bearing 

oblong-ellipsoid macroconidia 6-10x2-3 /xm (immersed, inconspicuous pycnidia containing 

mesoconidia 4-5-3x1-2-1-5 /am may also be present). All parts C- (t.l.c: nil). On 

lignum or decaying bark of old stumps 1. M. adnata (p. 108) 

lib Sporodochia absent . Thallus and/or pycnidia C -I- red (gyrophoric acid) or C - (t . 1 . c . : 'prasina 

unknowns') 12 

12a (lib) Thallus of pale to dark green (sometimes K-l- violet) goniocysts 12-60 /x,m; pycnidia, if 

present, 30-80 yam diam containing either (5-)5 -5-8x0-7-1 /xm microconidia, or 

(3-5-)4-6 X 1-2-1-7 ^tm mesoconidia. All parts C- (t.l.c. : 'prasina unknowns') 

34. M. prasina (p. 173) 

12b Thallus areolate, or scurfy (invaded by dematiaceous hyphae and foreign algae), or ± 

endoxylic; thallus and/or pycnidia usually C-l- red (gyrophoric acid) ; sometimes with curved 

or flexuose macroconidia 13 

13a (12b) With numerous pycnidia containing mesoconidia, 3-5-4-5x1-3-2 /xm, often 

extruded as conspicuous white blobs; in addition, pycnidia containing microconidia 

(4-)4-5-6(-6-5)x 0-7-1 /xm, or curved macroconidia 12-12x1 /xm, sometimes present; 

pycnidia walls with pale olivaceous pigment, K-l- violet. Usually on lignum, commonly on 

worked timber 11. M. denigrata(p. 127) 

13b Pycnidia usually with longer, curved or flexuose macroconidia; smaller pycnidia with microconidia 

sometimes present; mesoconidia unknown; pycnidial walls hyaline or greenish, K— 

On various substrata, rarely on worked timber 14 

14a (13b) Macroconidia curved or hamate, 21-40x1-1-5 /xm. Microconidia (5-)6-7(-7-7)x 

0-4-0-7 IX,. On various substrata but most commonly encountered without apothecia on 

bark of old trees (especially Quercus) in old woodlands 33. M. peliocarpa (p. 169) 

14b Macroconidia flexuose, 50-110x1 ixm. Microconidia (3-8-)4-5x0-5-0-7 /xm. Seen without 

apothecia on shaded lignum in woodland, and over bryophytes in open montane situations 

1. Micarea adnata Coppins, sp. nov. 

(Figs 7A, 36-37; Map 1) 

The species 

7. M. cinerea (p. 121) 

Thallus effusus, tenuis, ± laevis vel furfuraceo-granulosus, pallide griseo-viridis, plerumque nonnihil 

cereus. Algae cellulis 4—7 fxm diam. Apothecia pallide eburnea vel straminea vel demum pallide rufescen- 

tia, immarginata, adnata, primum planiuscula mox convexa, 0-2-0-4(-0-6) mm diam, interdum coalescentia 

in fasciculosos noduliformes ad 2 mm diam. Hymenium 35-40 îm altum, ± hyalinum. Ascosporae 

ellipsoideae, ovoideae, oblongo-ellipsoideae, oblongo-ovoideae vel oblongae, (O-)l-septatae, 9-16x 

.


3-5 />tm. Paraphyses numerosae, ramosae et anastomosantes, c. 1-1-5 jxm latae, apicibus vix incrassatis. 

Hypothecium hyalinum. Excipulum bene evolutum et manifestum praesertim in apotheciis junioribus. 

Conidiomata pycnidiiformia et sporodochiiformia. Pycnidia pauca et inconspicua, ± immersa, alba, c. 

40-60 /xm diam, producentia conidiis cylindricis 4-5-3x 1-2-1-5 ^tm. Sporodochia plerumque numerosa et 

conspicua, alba, pulvinata, 20-250 jxm diam, producentia conidiis cylindricis vel oblongo-ellipsoideis 

6-5-9-5X 2-3-3 ^tm. Thallus et apothecia, K— , C— 

, PD-; sine materia chemica. 

Typus: Caledonia, Argyll, Dunoon, Benmore, ad River Eachaig, ad corticem Alni, 18 xi 1977, leg. B. J. 

Coppins 3256 (E-holotypus). 

Thallus effuse, thin, ± smooth or finely scurfy-granular, pale grey-green, rather waxy in 

appearance; white, arachnoid, prothalline hyphae sometimes visible (x50). Phycobiont 

micareoid, cells 4-7 jxm diam. 

Apothecia usually numerous but occasionally few or absent, pallid to pale straw-brown or pale 

reddish brown; at first ± plane, adnate, ± immarginate but often with a white rim, c. 30-50 /am 

wide, from which a few white, arachnoid hyphae are sometimes seen to arise and penetrate into 

the thallus; later becoming convex although remaining adnate and never becoming constricted 

below; dispersed, or often confluent, and frequently coalescing to form large, irregular, waxy, 

nodulose clusters; individual apothecia 0-2-0-4(-0-6) mm diam, clusters up to 2 mm broad. 

Hymenium 35-40 /xm tall, hyaHne or dilute straw-yellow, K-. Asci clavate, 30-35x10-12 /xm. 

Spores ellipsoid, ovoid, oblong-ellipsoid, oblong-ovoid or oblong, often slightly constricted at 

the septum, (0-)l -septate, 9-16x3-5 fxm. Paraphyses numerous, branched and anastomosing, 

c. 1-1-5 fjLm wide and scarcely widening above; apices much branched and entangled. Hypothecium 

c. 40-120 /xm tall, hyaline; hyphae hyaline, c. 1-2 ^tm wide, interwoven but becoming ± 

vertically orientated towards the hymenium, intermixed with short-celled ascogenous hyphae 

that are up to 5 /xm wide. Excipulum well developed and clearly seen in young apothecia, c. 

30-35 /xm wide; hyphae radiating, branched and anastomozing, c. 1-2-5 /xm wide. 

Conidiomata of two types, one pycnidial, the other sporodochial. Pycnidia sometimes present 

but then usually few in number and inconspicuous, white, immersed, c. 40-60 îm diam; 

conidiogenous cells ± cylindrical, 6-10x1-1-5 /xm; conidia (mesoconidia) cylindrical, sometimes 

faintly biguttulate, 4—5-3xl-2-l-5 /xm. Sporodochia usually present, numerous and 

conspicuous, resembling small apothecia, white or pallid, pulvinate, 80-250 /xm diam; conidiogenous 

cells cylindrical, 10-20x1-7-2 fxm; conidia (macroconidia) cylindrical to oblong 

ellipsoid or occasionally oblong-obovoid, simple, eguttulate, 6-5-9-5x2-3-3 fxm. 

Chemistry: Thallus and apothecia K— , C— , PD-; no substances detected by t.l.c. 

Observations: Micarea adnata is easily recognised by the combination of a pale grey-green, ± 

waxy thallus, pale, adnate to clustered apothecia and, above all, by the presence of sporodochia. 

These cushion-like conidiomata, which resemble small apothecia, produce conidia externally 

over their surface; such structures are unknown in any other Micarea, and appear not to have 

been previously reported for lichenised fungi. The rather large, non-septate, oblong conidia 

produced by the sporodochia are probably homologous to the curved or filiform and often 

septate macroconidia of such species as M. cinerea, M. denigrata, and M. peliocarpa. Some 

specimens of M. adnata have a few pycnidia with typical mesoconidia, but microconidia have not 

yet been found. 

It is curious that this most distinctive species had not previously been afforded taxonomic 

recognition, although it was well known by Arnold who erroneously referred it to Biatorina 

prasiniza, a synonym of M. prasina. It is with forms of M. prasina with pallid apothecia that M. 

adnata is most likely to be confused. However, the former has a non-waxy, granular thallus 

composed of goniocysts, apothecia which soon become constricted below, usually smaller 

spores and an apparent absence of macroconidia. M. prasina is sometimes found to have 

pycnidia containing mesoconidia similar to those of M. adnata, but more frequently it has 

pycnidia containing microconidia. In addition, well developed specimens oiM. prasina have one 

of three distinctive substances detectable by t.l.c, whereas M. adnata contains no lichen 

substances. M. anterior is similar to M. adnata in being lignicolous with pale apothecia and 

spores that are mainly 1-septate; however, its apothecia are usually darker, varying from a pale


reddish or orange-brown to red-brown, with spores that are relatively narrower and ± fusiform, 

and a reddish brown excipulum. Furthermore, M. anterior has its mesoconidia borne in reddish 

brown, stalked pycnidia. 

Habitat and distribution: M. adnata is usually found on old stumps and associated decaying 

bryophyte mats and peaty debris; the Ugnum of such stumps is usually soft and riddled with holes 

bored by insects. In addition, it is found on the loose bark of trunks of old trees, including ^/nwj^, 

Betula, Corylus, Quercus, Abies, and Pinus. It usually occurs in extensive patches with few 

associated lichens, but amongst those noted are Cladonia spp. (including C. coniocraea and C. 

squamosa), Micarea prasina, Parmelia laevigata, and Platismatia glauca. Most collections are 

from old, ± natural deciduous or coniferous woodlands, but in western Scotland it is found also 

in old estate woodlands (possibly created from former ± natural woodland) containing many 

exotic trees. 

The distribution of M. adnata in Britain is centred on western Scotland, with a few outlying 

localities in Cumbria and north Wales. It is so far unrecorded for south-west England or western 

Ireland, and it should be sought for in the old oak-woods of those regions. Its distribution 

coincides with an annual rainfall of over 1000 mm per year, distributed over at least 160 rain 

days, and is attributable to the General Western group of Coppins (1976). Similarly, many of its 

localities on mainland Europe are in regions with a high annual precipitation (over 1000 mm per 

year), viz. southern Bavaria, Austria (Steiermark), and Switzerland (Bern). However, there are 

a few specimens from lower rainfall districts, namely, middle and northern Bavaria, and eastern 

Sweden (Sodermanland), perhaps in situations where local topographical or vegetational 

features provide a microclimate sufficient to compensate for the low rainfall. If the distribution 

of M. adnata is largely controlled by microclimatic conditions, usually resulting from a high 

rainfall, then it should be expected to extend to Norway, at least the south-western part of that 

country. 

Exsiccata: Arnold Lich. Mon. 244 (BM ex K, M). Britzelmayer Lich. Exs. 174 (M). 

2. Micarea alabastrites (Nyl.) Coppins 

(Figs 2, 6, 8; Map 2) 

in Topham & Walker in Lichenologist 14: 66 (1982). - 

Lecidea alabastrites Nyl. in Flora, Jena 62: 207 (1879). - Bilimbia sphaeroides var. alabastrites (Nyl.) A.L. 

Sm., Monogr. Br. Lich. 2: 138 (1911). Type: Ireland, West Galway, Derryclare, near Kylemore, on 

bryophytes on bark, 1878, C. Larbalestier (H-NYL 18656-hoIotype!; BM-isotype!). 

Thallus effuse, sometimes partly immersed in the substratum (especially when on lignum), 

but usually well developed on the surface of the substratum as crowded, confluent, ± 

convex-hemispherical to subglobose areolae. Areolae dull whitish, greenish white, or pale dull 

green, matt or slightly glossy, c. 40-140 /x,m diam, or up to 250 îm if containing a pycnidium of 

the macroconidial anamorph. Areolae in section, ecorticate but with a hyaline amorphous 

covering layer c. 4-10 jxm thick. 

Apothecia usually numerous and crowded, sometimes confluent and coalescing in groups of 

up to eight apothecia, adnate, plane to shallow-convex or convex-hemispherical, occasionally 

tuberculate, large, shallow-convex apothecia sometimes flexuose, often with an indistinct, 

slightly paler margin that is flush with the level of the disc, whitish, cream-white, ivory-white or 

pallid, 0-2-0-7 mm diam, or up to 0-9 mm diam when tuberculate. Hymenium 45-55 yam tall, 

hyaline, or dilute straw in upper part. Asci clavate, 42-50x12-16 /xm. Spores fusiform or 

clavate-fusiform, sometimes slightly curved, 3(-5)-septate, occasional 7-septate spores encountered, 

(16-)18-26(-29)x(4-)4-5-5(-6) jxm. Paraphyses numerous, branched, often anastomosing, 

c. 1-1-5 /xm wide; apices often more richly branched and entangled, often slightly 

incrassate to T8 /xm. Hypothecium c. 45-100 /am tafl; hyphae interwoven, c. 1-T7 /xm; 

ascogenous hyphae with swollen cells c. 2-4 /xm wide. Excipulum well developed (laterally c. 45 

/x wide in young apothecia), hyaline, of richly branched and anastomosing, radiating hyphae, c. 

1-T5 /xm wide.


Map 1 Micarea adnata # 1950 onwards O Before 1950 

Pycnidia usually present, white with hyaline walls, of two types: (a) immersed in areolae, c. 

100-220 /xm diam, ostiole often widely gaping; conidia (macroconidia) markedly curved, often 

sigmoid, sometimes faintly 3-5-septate, 21-55 xc. 1 (xm; (b) ± sessile, c. 50-100 ^tm diam, 

ostiole not, or only slightly gaping; conidia (microconidia) narrowly fusiform-cylindrical, 

5-7x0-5-0-7/am. 

Chemistry: Thallus and apothecia C-l- red; t.l.c: gyrophoric acid. 

Observations: M. alabastrites is morphologically and chemically very similar to M. peliocarpa, 

and it could easily be dismissed as representing a pale, shade form of that species. Apart from the 

complete absence of pigment from all its tissues, M. alabastrites is subtly different from M. 

peliocarpa in having slightly larger apothecia and spores. Shade forms of M. peliocarpa are 

usually sparingly fertile with scattered apothecia, whereas the apothecia of M. alabastrites are 

almost invariably numerous, crowded and often confluent. My early doubts regarding the 

distinction of these two species were dispelled by the collection, on several occasions, of M. 

alabastrites with adjacent thalli of M. peliocarpa with blackish apothecia; the slight differences in 

the sizes of apothecia and spores were confirmed with these collections. The two species also 

differ somewhat in distribution and habitat; M. peliocarpa is widely distributed throughout 

much of Europe on a wide range of substrata (tree trunks, mossy rocks peaty soil etc.); whereas 

M. alabastrites has a hyperoceanic distribution and occurs on tree trunks (or on bryophytes 

thereon) and occasionally lignum, but apparently never on mossy rocks or peaty soil. 

In the recent checklist of British lichens (Hawksworth et al, 1980) I mistakenly placed


Table 4 Diagnostic features for the separation of Micarea alabastrites, M. cinerea, and M. peliocarpa. 

Apothecia size (mm) 

Apothecia colour 

Hymenium height (/xm) 

Spore septation 

Spore length (/xm) 

Spore breadth 

Macroconidia shape 

Macroconidia length (/xm) 

Thallus colour 

peliocarpa alabastrites cinerea 

0.14^0-4(-0-6) 

pallid to black 

40-55 

(l-)3(-5) 

(ll-)15-23(-24) 

3-5(-6) 

curved-sigmoid 

21^0(-50) 

greenish white to 

dark blue-grey 

0-2-0-7 

whitish or pallid 

45-55 

3(-7) 

(16-)18-26(-29) 

(4-)4-5-5(-6) 

curved-sigmoid 

21-55 

greenish white or 

pale green 

0-2-0-7 

pallid to black 

55-70 

(3-)7 

(19-)23-34(-38) 

4-5-6 

flexuose 

50-110 

greenish white to 

dark blue-grey 

'alabastrites' as a synonym of M. cinerea. At that time I had only seen the fragmentary isotype of 

Lecidea alabastrites in BM; I found this to have a few 7-septate spores and considered it a 

juvenile, shade-form of M. cinerea. Subsequent examination of the holotype (a large, healthy 

specimen) proved me to be wrong, and that it actually belonged to a hyperoceanic species which 

I had intended to describe as new to science. M. cinerea is another close relative of M. 

alabastrites and is more prone to occur with ± pallid apothecia than M. peliocarpa. It differs 

from both these species in having spores which are mostly 7-septate at maturity, and macroconi- 

dia which are flexuose and much longer; for a comparison of these three species see Table 4. 

Occasional 7-septate spores have been found in several specimens of M. alabastrites, but they 

never number more than two or three in each squash preparation. 

M. alabastrites is one of the several lichens, named 'Bacidia (or Bilimbia) sphaeroides' by 

British lichenologists. This name is based on Lichen sphaeroides Dickson, which is a species of 

Biatora Fr. (non Ach.) and currently known as Catillaria sphaeroides (Dickson) Schuler. The 

name Bacidia sphaeroides as commonly used by Scandinavian workers refers to a species (not 

known in Britain) which also belongs in Biatora; its correct name (basionym) is probably 

Bilimbia tetramera de Not. 

Habitat and distribution: In the British Isles M. alabastrites is mainly found in communities of 

the Parmelietum laevigatae association on the trunks of Quercus, Betula, and, more rarely, 

Alnus, Fagus, Crataegus, Ilex, Pinus, Pseudotsuga, and Juniperus. Associated lichens include 

Bryoria fuscescens, Catillaria pulverea, Cladonia spp., Hypogymnia physodes, Lepraria incana 

agg., Micarea cinerea, M. peliocarpa, M. stipitata, M. synotheoides, Mycoblastus sanguinarius, 

M. sterilis, Parmelia crinita, P. laevigata, P. saxatilis, Platismatia glauca, Sphaerophorus 

globosus, and Usnea spp. Occasionally it is found on lignum of fallen decorticate trunks, 

especially in the western native pinewoods; associated species include Lecidea granulosa agg., 

Micarea lignaria, M. peliocarpa, and Platismatia glauca. To date, it is not known to occur on 

mossy boulders, directly on rock, or on the ground on peaty debris, etc. 

Its distribution in the British Isles is correlated with areas experiencing at least 180 'wet days' 

per annum and is referable to the 'General Western Group' of Coppins (1976). Elsewhere it is 

known from western Norway (Hordaland), the Azores (on Cryptomeria), and the Canary 

Islands {on Erica arborea). 

3. Micarea anterior (Nyl.) Hedl. 

(Figs7B,38C-D) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 76, 86 (1892). - Lecidea anteriorNyl. in Flora, Jena 

58: 299 (1875). - Catillaria anterior (Nyl.) Zahlbr., Cat. lich. univ. 4: 29 (1926). Type: Finland, Tavastia 

australis, Asikkala, 1863, /. P. Norrlin (H-NYL 21655 - lectotype!; H - isolectotypes!). 

Micarea anterior i. diluta Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 76, 86 (1892). Type: 

Sweden, Halsingland, Jarvso, vii 1890,7. T. Hedlund{S-ho\o\ypt\; UPS -isotype!).


Map 2 Micarea alabastrites # 1950 onwards O Before 1950 

Thallus effuse, endoxylic, or epixylic, then whitish and irregularly verrucose to 100 /xm thick, 

sometimes becoming secondarily rimose, but without distinct primary areolae; in section, 

without an amorphous covering layer. Phycobiont micareoid, cells 4-7 ^tm diam. 

Apothecia numerous, few, or absent, immarginate, convex, adnate, often becoming tuberculate, 

pale reddish brown, dull orange-brown or red-brown, 0- 15-0-35 mm diam, or to 0-6 mm 

when tuberculate, Hymenium 38-50 /zm tall, ± hyahne with irregular, dilute red-brown (K— , or 

dulling) blotches, especially in upper part. Asci clavate or cylindrical-clavate, 35^5 x 10-12 /xm. 

Spores oblong-ovoid to fusiform or ± bifusiform, often slightly curved, (0-)l(-2 or ?-3)-septate, 

9-14(-16)x2-5-4(-4-5) jxm. Paraphyses rather numerous, simple or sparingly branched, many 

becoming much branched above with their apices entangled, 0-6-1 /xm wide; apices often 

irregularly incrassate, hyaline and up to l-5(-2) îm wide, or sometimes thickened with reddish 

brown pigment and up to 3 ^tm wide. Hypothecium c. 60-110 ^lm. tall, hyaline. Excipulum thin 

but clearly seen in sections as a reddish brown reflexed zone; hyphae radiating, branched and 

anastomosing, hyaline and c. 1 fxm wide, but becoming thickened with brownish pigment and up 

to 2 jxm wide in the outer excipulum. 

Pycnidia usually numerous; of two types: (a) sessile on old apothecia, but more usually borne 

on slender stalks (pycnidiophores) which arise from the thallus or in some cases from old 

apothecia; 70-250 yam tall and 40-70 ^tm diam (overall); stalks pallid but often with reddish 

brown blotches, glabrous, simple, or sometimes branched and bearing two or three pycnidia; 

pycnidia (excluding stalk) short cylindrical or slightly tapering inwards above, 40-80 /xm tall and 

40-70 /xm diam, with dark reddish brown walls, K- or duUing; conidia {mesoconidia) cylindric-


al, often faintly biguttulate and slightly constricted in the middle, 3-5-4-5xl-2-l-6 /xm; (b) 

sessile on the thallus or old apothecia, c. 30-40 /xm diam, with dark reddish brown walls, K— or 

dulling; conidia (microconidia) narrowly cylindrical, (3-2-)3-6-4-5x0-7-0-9 /xm. 

Chemistry: Thallus K— , C— 

detected by t. I.e. 

, PD— 

; sections of thallus and apothecia C-; no substances 

Observations: Micarea anterior is characterised by its reddish brown apothecia, rather 

narrow, mostly 1-septate spores, pale pycnidiophores bearing dark brown pycnidia, and a ± 

endoxylic, or thin and irregularly verrucose, thallus. Stalked pycnidia are a feature of several 

other hgnicolous species (e.g. M. melaeniza, M. misella, M. nigella, and the occasionally 

lignicolous M. botryaides), but in those species the pycnidiophores appear black and have 

different pigmentations when examined microscopically. The pycnidiophores of M. hedlundii 

are brownish but distinctly tomentose, and those of the normally corticolous M. pycnidiophora 

and M. stipitata are entirely hyaline. M. adnata has its mesoconidia produced in immersed 

pycnidia and has macroconidia borne on small apothecia-like sporodochia; further distinguishing 

features from M. anterior include a ± waxy appearance of the whole plant, mostly 

paler apothecia, and relatively broader spores with rounded apices. Richly fertile specimens of 

M. anterior could be confused with Catillaria erysiboides {q. v. ) but that species has marginate 

young apothecia, short ovoid spores, and a non-micareoid phycobiont. 

Habitat and distribution: M. anterior is found on the rather soft and smooth lignum of 

decorticate trunks of Picea and Pinus, and associated with such species as Cetraria pinastri, 

Cladonia spp., Hypogymnia physodes, Lecidea icmalea, Lepraria incana agg., Micarea adnata, 

M. contexta, M. misella, M. prasina, Parmeliopsis ambigua, P. hyperopta, Pertusaria pupillaris, 

and the non-lichenised Cryptodiscus pallidas. It is a rare or overlooked species, apparently 

confined to Scandinavia, from where it is known from several locahties in middle and southern 

Sweden, and south-west Finland. 

Exsiccata: Malme Lich. Suec. 22 (M, S). 

4. M. assimilata (Nyl.) Coppins, comb. nov. 

(Fig. 9; Map 3) 

Lecidea assimilata Nyl., Lich Scand.: 221 (1861). Type: Norway, Nordland, Helgeland, M. N. Blytt 

(H-NYL 16556- lectotype!). 

Lecidea assimilata a. [var.] irrubata Th. Fr., Lich. Scand. 2: 522 (1874); nom. inval. (Art. 26). 

Thallus growing on bryophytes, plant debris or sandy soil, composed of confluent, irregular, 

convex-verrucose, sometimes flattened and subeffigurate areolae that are often intermixed with 

cephalodia. Areolae white or brownish white, matt or sometimes glossy, 0-08-0-4 mm diam; in 

section, sometimes with a hyaline amorphous covering layer up to 7 /xm thick, outermost hyphae 

hyahne; a ± algal-free medulla sometimes differentiated in large areolae. Thallus hyphae c. 

T8-3 /xm diam. Phycobiont micareoid, cells 4—7 fxm diam. Cephalodia often present, irregularly 

globose and hidden amongst the areolae, sometimes visible externally as brown areolae-like 

structures, 0-2-0-4 mm diam; containing Nostoc, cells 3-5 fxm diam. Less often present are 

irregular, rather loose clusters (? cephalodia) of Stigonema. 

Apothecia numerous, immarginate, convex to subglobose, matt or ± glossy (subnitid), 

0-3-0-8 mm diam, sometimes forming tuberculate clusters up to T5 mm diam. Hymenium, 

45-50 /xm tall; upper part (epithecium) dark aeruginose, olivaceous or brownish-green, K— 

HNO3+ red; mid-hymenium dilute greenish; lower hymenium dilute greenish and K— , or dilute 

purplish and K-l- purple intensifying or K-l- sordid green. Asci clavate, 45-50x 12-14 /xm. Spores 

oblong-ellipsoid to oblong-fusiform, 0(-l)-septate, (10-)12-16(-19)x3-5 /xm. Paraphyses 

numerous, simple below, but often forked or branched above, 1-5-1 •7(-2) jxm wide, sometimes 

widening above to 3 /xm; apical walls hyaline although surrounded by densely pigmented 

gel-matrix. Hypothecium c. 150-400 /xm tall, dark purple brown, K-t- purple intensifying or 

(especially in upper part) K-l- dark green; all parts HNO3-I- purple-red; hyphae interwoven but 

,


± vertically orientated near the hymenium, c. 1-7-2-7 fxm wide, embedded in a densely 

pigmented matrix; ascogenous hyphae c. 1-7-2-7 /xm wide, embedded in a densely pigmented 

matrix; ascogenous hyphae c. 2-5-5 fxm wide. Excipulum reflexed, mottled reddish brown or 

sordid olivaceous; hyphae radiating, branched and anastomosing, 1-5-2 /xm wide. 

Pycnidia rare, semi-immersed to sessile, black, 40-100 /^m diam; walls dull reddish brown, or 

dull olivaceous in part; conidia (? microconidia) cylindrical, 6-9x1-1-5 /xm. 

Chemistry: Thallus K— , C— , KC— 

, PD — ; t.l.c: no substances. 

Observations: Micarea assimilata is characterized by its conspicuous, whitish, verrucose 

areolae which are often intermixed with brown cephalodia, rather large, convex, black 

apothecia, green (K— ) epithecium, dark purple-brown hypothecium and predominantly simple, 

large spores. It is apparently closely related to M. crassipes, M. incrassata, M. melaenida, and M. 

subviolascens , all of which share rather stout, simple or sparingly branched paraphyses, rather 

stout excipular hyphae, dark hypothecia and an absence of lichen substances. In many ways M. 

assimilata is identical to M. incrassata and both occur in similar habitats; indeed the latter has 

often been considered a variety (var. infuscata) of the former (e.g. Anderson, 1964; Hertel, 

1977). However, M. incrassata differs in several respects, the most important of which is its quite 

different, red-brown hypothecium which lacks any trace of purple pigmentation in water, K or 

HNO3. In addition, it has an often darker thallus, less prominent apothecia and a generally 

higher proportion of septate spores. M. subviolascens is similar in appearance to M. assimilata, 

but grows on rock and has a green, K-l- violet epithecial pigment. M. melaenida differs from the 

last three species in its comparatively small apothecia and less conspicuous thallus, absence of 

green pigmentation in apothecial tissues, preponderance of 1-septate spores, shorter conidia, 

and confinement to argillaceous, or fine-grained, mineral soils. M. crassipes occurs in much the 

same habitats as M. assimilata and the two species have been much confused. However, M. 

crassipes is easily distinguished by its apothecia which are thinly marginate (at least when young) 

and markedly constricted below, often being turbinate or even short-stipitate; also by its thallus 

which is composed of small granular-areolae which sometimes proliferate to give it an isidiose 

appearance. In section, the apothecia of M. crassipes display a well-developed excipulum and a 

distinctly two-zoned hypothecium. 

Pycnidia are rare and difficult to find in members of the M. assimilata group, and have not yet 

been found at all in M. subviolascens. The remaining species appear to have one conidium type 

only; although the conidia are rather large, they afe probably microconidia. However, the 

relatively shorter and broader conidia of M. crassipes may be better described as mesoconidia. 

Cephalodia are here reported from M. assimilata and M. incrassata, apparently for the first 

time. In both cases the blue-green alga concerned is Nostoc, which loses its filamentous form. In 

addition, irregular clusters of partially disrupted Stigonema filaments are sometimes present 

amongst the areolae of M. assimilata, M. incrassata, and also M. subviolascens. Hyphae, 

presumably belonging to the lichens, are seen to ramify through these clusters; but whether or 

not these loosely organised structures can be considered to be cephalodia is a problem requiring 

further detailed anatomical and experimental investigation. Cephalodia or cephalodia-like 

structures have not been encountered in any other Micarea species. 

The closely related Lecidea limosa Ach. and L. stenotera (Nyl.) Nyl. have often been confused 

with M. assimilata and M. incrassata, but they can be distinguished by their pale hypothecia, 

paraphyses which are individually supplied with a dense, hyaline, gelatinous sheath, and more 

organised excipular structure. Together with Catillaria contristans (Nyl.) Zahlbr. they probably 

represent a distinct genus within the Lecideaceae, although perhaps not far removed from 

Micarea. In British herbaria (at least) there has been much confusion between M. assimilata and 

Lecidea hypnorum Lib. The latter has marginate, brown-black apothecia, simple paraphyses, 

0(-3)-septate spores which become straw coloured and finely warted with age, a reddish brown 

excipulum composed of radiating, stout, pachydermatous, heavily conglutinated hyphae, c. 

3-5 /xm wide, and bears little resemblance to a Micarea. In addition, the hymenium, hypothecium 

and excipulum of L. hypnorum (and the related L. berengeriana (Massal.) Th. Fr. , and L. 

sanguineoatra auct.) often contain minute granules of a dark violet (K-l- aeruginose) pigment.


More understandable cases of mistaken identity involve Lecidea caesioatra Schaerer, which 

bears a close superficial likeness to M. incrassata. It can be distinguished from this Micarea by its 

dark blue-grey apothecia which have a caesious bloom when wet, excipulum of conglutinated (in 

K) hyphae, and non-micareoid phycobiont with cells c. 8-12 /xm diam. 

Habitat and distribution: M. assimilata grows on decaying bryophytes and plant detritus on the 

ground or on soil accumulation in rock crevices in arctic (i.e. low altitudes at high latitudes) or 

montane situations. The realisation that this species belongs in Micarea came so late in the 

present study, that I have been able to examine only a limited amount of herbarium material. To 

date, I can confirm its presence in northern Scandinavia and the single outlying locality in 

Scotland. Specimens purporting to be Lecidea assimilata from the Alps and other central 

European mountains have proved to be M. crassipes, M. incrassata, or 'Lecidea' species not 

referable to Micarea. Material of 'Lecidea assimilata'' from Greenland distributed as Hansen 

Lich. Greenland Exs. Ill and 247 belongs to Lecidea stenotera and L. limosa respectively. 

The Scottish locality is Ben Lawers in Perthshire, a mountain long renowned for its 

arctic-alpine flora (Anon, 1972; James, 19656). M. assimilata was collected there by W. L. 

Lindsay in 1856. Recent collections from Ben Lawers, and other Scottish localities, are referable 

to other species, such as Lecidea caesioatra, L. hypnorum, L. limosa, and Micarea incrassata. 

Exsiccata: Krypt. Exs. Vindob. 2268 (BM, H). Malme Lich. Suec. 216 (H). 

Map 3 Micarea assimilata ® Before 1950 + Micarea incrassata # 1950 onwards

5. Micarea bauschiana (Korber) V. Wirth & Vezda 

(Figs5,10A,40D;Map4). 


in Vezda & V. Wirth in Folia geobot. phytotax., Praha 11: 95 (1976). _ Biatora bauschiana Korber, 

Parerga lich.: 157 (1860). - Lecidea bauschiana (Korber) Lettau in Hedwigia 55: 28 (1914). Type: 

Germany, Baden-Wiirttemberg, Baden-Baden, on the way to Yburg, on porphyry, 1859, Bausch, 

Rabenh. Lich. Eur. 648 (M- lectotype!, sel. V. Wirth & Vezda (loc. cit); M-isolectotype!); additional 

isotype material distributed as Arnold Lich. Exs. 120 (BM!, M!). 

Lecidea infidula Nyl. in Flora, Jena 51: 475 (1868). Type: Jersey, C. Larbalestier (H-NYL p.m. 5413 - 

holotype!). 

Lecidea lynceola Th. Fr., Lich. Scand. 2: 561 (1874). Type: Norway, Christiana [Oslo], Tveten, 20 v 1868, 

N. G. Moe 257 (UPS -holotype!). 

Lecidea semipallens Nyl. in Flora, Jena 59: 234 (1876). Type: Ireland, West Galway, Lough Inagh, 1875, C. 

Larbalestier (H-NYL 19399 - lectotype!; isolectotypes: BM ex K!, H-NYL 19402!). 

Lecidea dilutiuscula Nyl. in Flora, Jena 59: 308 (1876). Type: England, South Devon, near Buckfastleigh, 

H. B. /fo//(BM- lectotype!; H-NYL 10754- isolectotype!). 

Lecidea rusticella Nyl. in Flora, Jena 61: 245 (1878). Type: Ireland, West Galway, Connemara, Tullywee 

Bridge, 1876, C. Larbalestier (H-NYL 20206- holotype!; topotypes ['1878']: BM!, BM ex K!). 

?Lecidea callicarpa Larbal. ex Leighton, Lich. Fl. Br., ed. 3: 266 (1879). Type: Ireland, West Galway, 

Glencorbot near Kylemore, 1877, C. Larbalestier (not seen; possibly in BM (incl. BM ex K) but not 

traced). 

?Lecidea semipallens var. obscurior Lang ex Havaas in Bergens Mus. Arb. 1935(2): 27 (1935). Type: 

Norway, Dalsb0, 1903, /. J. Havaas (not seen; not traced in BG or H). 

?Catillaria microsporaMas\o\a.in Ukr. bot. Zh. 30(5): 665 (1973). Type: USSR, Regio Volhyniensis, distr. 

Ljuboml., in pineto baud procul pag. Kamenca, ad saxa granitica, 11 vi 1969, W. R. Maslova (KWholotype, 

not seen; but a slide prepared from the holotype by H. Kilias examined). 

Thallus thin (


often with a swollen base up to 3 /xm wide; conidia (microconidia) cylindrical, eguttulate, 

4-6x0-8-1 fxm. A few collections with pallid apothecia have associated pycnidia, c. 60-100 /xm 

diam, with hyaline walls and containing (often biguttulate) mesoconidia 2-8-4-3X 1-1-3 /xm (see 

'Observations' below). 

Chemistry: All parts K- , C- , KC— , PD- ; no substances detected by t.l.c. 

Observations: Micarea bauschiana is closely related to, and often confused with, M. sylvicola 

(q.v.), and the two species often grow together. M. bauschiana is notoriously variable in the 

colour of its apothecia, but the full range of variation from pallid to blackish can be seen over 

small distances (a few centimetres) in a single population (Fig. 5: p. 30) and the variation appears 

to be solely phenotypic. 

When growing on iron-rich rocks the thallus of M. bauschiana is often oxydated. In a few 

specimens (including types of Lecidea rusticella) this oxydation has extended to the apothecia in 

which finely granular, ferruginous material has been deposited (especially in the epithecium and 

hypothecium); such specimens have been confused with M. lutulata and species of Protoblaste- 

nia. 

A few collections from Austria and Scotland (considered here as M. cf. bauschiana) have 

entirely pallid apothecia which are accompanied by pycnidia containing mesoconidia (see 

description above). Such pycnidia have not been detected in specimens of M. bauschiana with 

pigmented apothecia, and it is possible that the former collections represent a distinct taxon. 

However, one of these collections (Austria, Steiemark, Graz, Schockl N of St Radegund, 1978, 

Poeh (GZU) also has typical (for M. bauschiana) microconidia-containing pycnidia, and it may 

be that there is some environmental control (e.g. a response to very low light intensities) 

involved in the initiation of the mesoconidial anamorph. Further careful field observations and 

laboratory studies are required to establish the status of these seemingly anomalous collections. 

Forms of M. bauschiana (s. ampl. ) with pallid apothecia should be compared with the much 

rarer M. lithinella {q. v. ) which can be distinguished by its micareoid phycobiont. In addition, M. 

lithinella seems not to occur in the ombrophobous, aerohygrophilous Micareetum sylvicolae, 

and is a rather ombrophilous, substratohygrophilous species of damp, shaded rocks or stones. 

However, I must stress that this ecological interpretation of M. lithinella is based on the limited 

data available from the herbarium specimens and requires confirmation by field studies. 

Material recently distributed as 'M. bauschiana' in Hertel Lecid. exs. no. 54 is not this species. 

It has weakly marginate apothecia due to a well developed excipulum, and a large-celled 

(9-17 /xm diam) phycobiont, the cells of which are deeply penetrated by distinct haustoria (as in 

M. intrusa). It is probably an undescribed species of Micarea and will be treated in a later 

pubhcation following more critical studies. 

Habitat and distribution: M. bauschiana is a faithful member of the Micareetum sylvicolae and 

grows on rocks, stones, roots, and consolidated soil in dry underhangs in woodlands or sheltered 

valleys. In the British Isles M. bauschiana is the commonest member of the M. sylvicola group. It 

is particularly common in the north and west but also occurs in suitable situations (e.g. dry stones 

in sandy banks in woodlands) in the lowlands of south-west England. It seems to be widely 

distributed in Europe but I have not seen enough specimens to make an assessment of any 

distributional tendencies. It occurs in the Azores and the Canary Islands, but I have not seen any 

additional material from outside Europe. 

Exsiccata: Arnold Lick. Exs. 120 (BM, M), 1233 (M). Johnson Lick. Herb. AM p.p., 504 (HAMU). 

Larbal. Lick. Herb. 68 (BM), 305 p.p. (LD). Rabenh. Lich. Eur. 648 p. max. p. (M). Zwackh Lick. 

Exs. 279A-B, 594A-B, 595 (M). 

6. Micarea botryoides (Nyl.) Coppins 

(Figs 10b, 35, 38A; Map 5) 

in D. Hawksw., P. James & Coppins in Lichenologist 12: 107 (1980). - Lecidea apochroeella var. 

botryoides Nyl. in Flora, Jena 50: 373 (1867). - Lecidea botryoides (Nyl.) Zahlbr., Ca/. lich. univ. 3: 740


Map 4 Micarea bauschiana # 1950 onwards O Before 1950 

(1925). Type: Finland, Tavastia australis, Lammi, Evo, Lapinkallio, 1866, J. P. Norrlin 404 (H-NYL 

20685 p.p. -lectotype!; H-isolectotype!). 

Note: In the protologue Nylander states 'ad lignum putridum' but this is clearly an error (see Vainio 

1934: 361). The lectotype has been selected from a packet in the Nylander Herbarium containing two 

pieces of rock - one with M. lutulata, the other (lectotype) with the apothecia and characteristic stalked 

pycnidia of M. botryoides. It is possible that Nylander 's diagnosis was based in part on the apothecia of M. 

lutulata, which bear many similarities to those of M. botryoides. Because Art. 70 of the ICBN is no longer 

applicable Lecidea apochroeella var. botryoides cannot be rejected as being based on discordant elements; 

consequently the part containing M. lutulata is excluded, and that part including stalked pycnidia (as 

described in Nylander's diagnosis) is chosen as the lectotype. 

Thallus effuse, thin, more rarely developing into a thick, loose crust up to c. 0-4 mm thick, 

scurfy-granular, pale to dark dull green, dark olive-green, or dull greenish black, sometimes 

whitish buff when on dry, deeply shaded rocks. When very thin the thallus consists of flattened 

granules, c. 20-50 îm diam. , which are often dispersed and interconnected by white, arachnoid, 

prothalline hyphae. Thickening of the thallus is caused by the production of goniocysts (c. 16-30 

ixm diam), the first of which appear to arise by budding from the primary granules; when thick 

the thallus has a ± gelatinous appearance when wet. Phycobiont micareoid, cells 4—7 /xm diam. 

Apothecia usually few, more commonly absent, immarginate, at first convex-hemispherical, 

soon becoming ± globose and much constricted below (sometimes short-stipitate), often 

becoming tuberculate, black, or dark brown (in deep shade), matt, 0- 1-0-25 mm diam, or to 0-5 

mm when tuberculate. Hymenium 25-35 îm tall, hyaline or sometimes tinged dilute brown 

(K- , HNO3-) or dilute olivaceous (K- , or K± green intensifying, HNO3+ red) in places, but


always with numerous dark brown (K± olivaceous tinge, HN03± reddish tinge) vertical 

streaks. Asci cylindrical-clavate or clavate, 20-35x7-9 /xm. Spores ovoid, oblong-ellipsoid or 

oblong-ovoid, straight or slightly curved, 0-l(-3)-septate, 8-13(-16)x 2-3-3 •7(-4) /xm. Paraphyses 

rather scanty, of two types: p.p. evenly distributed, flexuose, simple or sparingly 

branched, sometimes anastomosing, thin, 0-7-1 fxm wide, sometimes widening above to 1 -7 /^m, 

walls hyaline throughout; p.p. fasciculate, simple or sparingly branched, stout, c. 2-2-5 /xm 

wide, sometimes widening above to 3-5 ixm, coated ± throughout by dark brown pigment. 

Hypothecium c. 60-120 /xm tall, dark reddish brown, K- or dulling, HNO3- or red tinge 

slightly intensifying; hyphae coated with dark brown pigment, c. 2-3 /xm wide, interwoven but 

becoming vertically orientated towards the hymenium and sometimes continuing into it as stout, 

fasciculate, pigmented paraphyses; ascogenous hyphae similarly pigmented, with short, swollen 

cells to 5 /xm wide. Excipulum indistinct, sometimes evident in young apothecia as a reflexed 

reddish brown zone concolorous with, or sHghtly paler than, the hypothecium; hyphae radiat- 

ing, branched and anastomosing, c. 1-1-5 /xm wide, their walls sometimes with a thin coating of 

brown pigment. 

Pycnidia always present and numerous, sessile or, more usually, distinctly stalked, black, 

50-400 /xm tall (including stalk) and 40-90 /xm diam; stalks simple, or branched and bearing up 

to six pycnidia; the 'stalk-part' below the current conidia-producing pycnidia often includes old 

pycnidia (see Fig. 35); the stalk and pycnidia are usually black but in extreme shade forms the 

stalk tissue may be ± colourless, contrasting with the dark brown or blackish, current and old 

pycnidia. In microscope preparations (at x400): pycnidiophore tissue dilute to dark fuscous or 

reddish brown, K- or dulling, HNO3- or red tinge slightly intensifying; pycnidial wall dark 

greenish brown, K- or K-l- green intensifying HN03-(- red. Conidiogenous cells ± cylindrical or 

elongate ampuUiform, often with swollen base which is thickened with brownish pigment, often 

with one or two percurrent proliferations, 3-5-7-5 x 1-1-4 /xm, base sometimes swollen to 2-5 /xm 

wide. Conidia (mesoconidia) ± cylindrical, often biguttulate, sometimes slightly constricted in 

the middle, 3-5-4-8x1-1-5 /xm. 


detected by t. I.e. 

, PD— 

; sections of apothecia and thallus C— ; no substances 

Observations: Micarea botryoides is easily recognised by its numerous, black, stalked 

pycnidia, whose walls are dark greenish brown (K- or K-l- green intensifying), and its normal 

occurrence on substrate other than hgnum. It is very similar to the rare M. melaeniza, but that 

species has shorter conidia, smaller, simple spores, and is apparently confined to lignum. M. 

misella often has stalked, black pycnidia, but is usually hgnicolous. However, it does occasionally 

occur on decaying bryophytes where it could be confused with M. botryoides. In such 

instances their separation is easy because the pycnidia of M. misella contain an olivaceous or dull 

brownish pigment that turns violet in K. When on rock the apothecia of M. botryoides could be 

confused with those of M. lutulata, but the latter species has smaller, simple spores, a 

non-micareoid phycobiont, and immersed pycnidia. When on soft lignum it should be compared 

with M. nigella which is superficially identical, but has a purple-brown (K-l- green) pigment in its 

apothecia and pycnidia. 

Sterile forms of M. botryoides have puzzled lichenologists for many years, being dismissed 

with such remarks as 'indeterminate pycnidia' or 'fungus'. In 1867 Leighton distributed sterile 

material of M. botryoides in his exsiccate (no. 388), as ' Lecidea sabuletorum var. milliaria (Fr.), 

spermagonia', evidently believing it to be the pycnidial state oi Micarea lignaria, the apothecia 

of which occur on some of his specimens . The 

identity of these pycnidia remained a mystery until 

my discovery of fertile material in Scotland in 1976 and my subsequent examination of the type 

material of Lecidea apochroeella var. botryoides in 1979. 

Habitat and distribution: M. botryoides is usually found as a constituent of the Micareetum 

sylvicolae in dry underhangs, growing on solid rock, loose stones, consolidated soil, tree roots, 

and loose mats ot moribund bryophytes; but it also occurs on rock, stones, and decaying 

bryophytes in wetter shaded situations. There are a few collections made from the soft.


Map 5 Micarea botryoides # 1950 onwards O Before 1950 

crumbling lignum of old stumps, but I have not seen it on the firmer, ± smooth lignum favoured 

by M. melaeniza and M. misella. In the environs of some industrial regions (e.g. the West 

Yorkshire conurbation) it has been found on bark at the bases of trunks oi Acer and Betula in 

sheltered woodlands. 

M. botryoides is a widespread species, especially in western and upland regions of Britain, and 

is certainly much more common than current records would suggest. Several of these records 

result from accidental gatherings, being subsequently identified on samples of other members of 

the Micareetum sylvicolae. This gives an indication as to how often this species must have been 

overlooked. It is little recorded outside Britain, and to date I know it only from Norway 

(Sor-Trondelag and Hordaland), Finland (Tavastia australis), and north-west France. 

Exsiccata: Leighton Lich. Brit. 388 (BM, BON, DBN, E, M). 

7. Micarea cinerea (Schaerer) Hedl. 

(Figs 11, 39; Map 6) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 81, 93 (1892). - Lecidea cinerea Schaerer, Lich. 

Heiv. spic, sect. 3: 156 (1826). - Bacidia cinerea (Schaerer) Trevisan in Linnaea 28: 293 (1856). - 

Bilimbia cinerea (Schaerer) Korber, Parerga Lich. : 

164 (1860). Type: Switzerland, 'ad infimos Abietum 

truncos, insylvaKonitz', L. E. Sc/zaerer (G-holotype!; M-? isotype!). 

Lecidea sphaeroides var. albella Schaerer, Lich. Helv. spic, sect. 4^5: 165 (1833). Type: Switzerland: 

'Schweiz' [on conifer bark], L. E. Schaerer (M - neotype!). See note (i). 

Biatora delicatula Korber, Denkschr. Feier ihres funfzigjah. Best, herausg. Schles. Gesellsch. voter. Kult.,


Breslau: 233 (1853). - Bilimbia delicatula (Korber) Korber, Syst. lich. Germ: 212 (1855). Type: lectotype 

as for Lecidea sphaeroides var. albella Schaerer. See note (ii) below. 

Bilimbia cinerea f. hypoleuca Stizenb. ex Arnold in Flora, Jena 58: 598 (1864). - Micarea cinerea f. 

hypoleuca (Stizenb. ex Arnold) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 82, 93 (1892). 

Type: Germany, Bayern, near Eichstatt, Affenthale, on young Picea abies in a wood, viii 1864, F. C. G. 

Arnold (M - lectotype!). 

Notes, (i) No material labelled Lecidea sphaeroides var. albella could be traced in G (Geissler, in litt.) 

and an undated specimen in M is selected as neotype. If it can be proven that this specimen was collected 

before 1833, it should be regarded as a lectotype. 

(ii) No material referred to in the protologue of B. delicatula has been traced in L, WRSL or elsewhere. 

However, Korber gives 'Lecidea sphaeroides a albella Schaer. Enum. 193.' as a synonym. This citation 

refers to Schaerer (1850) and is an indirect reference to Lecidea sphaeroides var. albella Schaerer (1833). 

Consequently I have typified both names with the same specimen in M. 

Thallus effuse, sometimes partly immersed in the substratum (especially when on lignum), 

more usually developed on the surface of the substratum as rounded, shallow-convex, hemispherical 

or ± globose areolae. Areolae scattered to ± contiguous, smooth, greenish white to 

blue-grey, or more rarely becoming dark grey, often dark coloured on upper surface but 

greenish white below, c. 40-160 fxva diam or up to 300 jxm diam if containing a pycnidium of the 

macroconidial anamorph. Areolae in section, ecorticate but with a hyaline amorphous covering 

layer c. 2-5 /xm thick; outermost hyphae often with grey-green to aeruginose walls, K-, 

HNO3-I- red. Phycobiont micareoid, cells 4-7 /xm diam. 

Apothecia usually numerous (see 'observations'), adnate, plane to convex, sometimes 

becoming tuberculate, sometimes with an indistinct margin that is ± flush with the level of the 

disc, pale leaden-grey to grey-black (margin often paler), or ivory-white or pallid in shade forms, 

(0-2)0-3-0-7 mm diam or up to 1-3 mm when tuberculate. Disc matt and finely roughened, but 

margin smooth and often ± glossy. Hymenium 55-70 /xm tall, hyaline, but usually olivaceous or 

aeruginose (K— , HNO3+ red) in upper part (epithecium). Asci clavate 50-65x15-20 /am. 

Spores fusiform, often slightly curved, (3-)5-7-septate, (19-)23-34(-38)x 4-5-6 ixm. Paraphyses 

numerous, branched and often anastomosing, 1-1-4 /xm wide; apices often more richly 

branched and entangled, often slightly incrassate to c. 1-8 îm, or 2-5 /xm due to thickening by 

greenish pigment. Hypothecium c. 40-70 /xm tall, hyaline; hyphae interwoven, c. 2-4 /xm wide. 

Excipulum well developed, hyaline or pale straw, or richly branched and anastomosing hyphae, 

c. 1-1-5 /xm. 

Pycnidia frequently present, of two types: (a) immersed in areolae, white or faintly greenish 

around the ostiole, 160-300 /xm diam, ostiole often widely gaping; conidia (macroconidia) ± 

straight or flexuose, filiform, 9-17-septate, 50-1 10 xc. 1 /xm; (b) ± sessile, white, c. 40-70 /xm 

diam, ostioles not, or only slightly, gaping; conidia (microconidia) narrowly fusiform-cylindrical 

(3-8-)^5xO-5-0-7/xm. 

Chemistry: Thallus and whitish apothecia K-, C-l- red, PD-; apothecia in section C-l- red; 

t.l.c: gyrophoric acid. 

Observations: M. cinerea is very variable with regard to the colour of its apothecia and thallus; 

as with M. peliocarpa this variation is due to the amount of green pigment produced in response 

to environmental factors, especially light. Morphologically and chemically M. cinerea is closely 

allied to M. alabastrites and M. peliocarpa; the diagnostic features of each of these three species 

are compared in Table 4. Forms of M. cinerea with blackish apothecia can be confused with M. 

lignaria (including var. endoleuca) but on close inspection the latter will be seen to differ in its ± 

globose apothecium with a poorly differentiated excipulum (in vertical section), usually 

brownish or greenish upper hypothecium, less strongly branched paraphyses and chemistry. 

M. cinerea is usually well fertile with numerous apothecia, but sterile forms with numerous 

macroconidia-containing pycnidia have been encountered; the macroconidia are about twice as 

long as those found in M. peliocarpa and M. alabastrites and so are readily identifiable (see 'Key 

to species without apothecia, pp. 107-108).


Habitat and distribution: In the British Isles M. cinerea is mostly found in communities of (or 

closely akin to) the Parmelietum laevigatae on the trunks or over bryophytes thereon of Quercus, 

Betula and, less often, Alnus, Corylus, Fraxinus, Ilex, Sorbus, Larix, Pinus, and Pseudotsuga. 

Associated species in these communities include Catillaria pulverea, Cladonia coniocraea, C. 

macilenta, C. squamosa, Haematomma caesium, H. elatinum, Lecidea icmalea, Lepraria incana, 

Micarea alabastrites, M. stipitata, M. synotheoides, Mycoblastus sterilis, Parmelia laevigata, P. 

saxatilis, Platismatia glauca, Stenocybe septata, Thelotrema lepadinum, Trapelia corticola ined. , 

Frullania tamarisci, Lejeunea ulicina, Scapania gracilis, Dicranum fuscescens, and Hypnum 

cupressiforme. Less commonly it is found in the Ugnum of fallen, decorticated trunks. Occurrences 

on other substrata are rare although I have found it on one occasion growing directly on 

rock (Coed Hafod in Denbigh) where it was associated with Trapelia involuta on the top of a 

dry-stone wall in an oak-birch wood. In addition, it was found on mosses on epidiorite at an 

altitude of 1000 m in the Ben Alder range of Inverness-shire by Dr O. L. Gilbert; the specimen is 

sterile but has pycnidia with the characteristic macroconidia. In Britain M. cinerea usually occurs 

at lower altitudes (mostly below 300 m), and exhibits a distribution pattern attributable to the 

General Western Group of Coppins (1976). However, there are two outlying easterly localities 

(in North Northumberland and East Perth) both of which are sheltered, more or less undisturbed, 

valley woodlands. 

From outside Britain I have seen material of M. cinerea from Hordaland and Rogaland in 

western Norway, the east Sudety and Vysoke Tatry of Czechoslovakia, southern Germany, and 

the Swiss, Austrian and Italian Alps. Most collections from southern Germany and the Alps are 

Map 6 Micarea cinerea • 1950 onwards O Before 1950


from the trunks of conifers (Picea, Pinus, and Abies), but they also included specimens from the 

trunks of A Inus and Betula, lignum of stumps, fallen trees, and old fence posts. In addition, it 

was collected several times on thin twigs of Picea by F. Arnold and his contemporaries, and it is 

possibly of interest to note that I do not know it as an inhabitant of twigs in north-west Europe. 

Unlike M. alabastrites and M. peliocarpa, M. cinerea is not known from Macaronesia, and I 

have not seen any material of it from outside Europe. 

Exsiccata: Arnold Lich. Exs. 548 (M), 549 (BM ex K, M). Arnold Lich. Mon. 47 (BM ex K), 1 15 (BM ex 

K, M, MANCH), 116 (BM ex K, M). Britz. Lich. Exs. 846 (M). Hepp Flecht. Eur. 21 p.p. (BM, M). Lojka 

Lich. Hung. 60 (M). Vezda Lich. Set. 1087 (BM, S). Zwackh Lich. Exs. 898 (M). 

8. MicareacontextaHedl. 

(Figs 12A, 40A-B) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 83, 96 (1892). - Catillaria contexta (Hedl.) Zahlbr, 

Cat. lich. univ. 4: 35 (1926). Type: Sweden, Halsingland, Ovanaker, 1891, J. T. Hedlund (S - 

lectotype!). 

Thallus effuse, endoxylic, inapparent or evident as a slight bleaching of the wood, consisting 

of scattered, small, rounded granules (c. 15-25 pm diam) buried between the wood fibres; 

external hyphae of granules with dark green walls, K— , HNO3-I- red. Phycobiont micareoid, 

cells 4-7 pm diam. 

Apothecia numerous, immarginate, ± globose from the start, often becoming tuberculate, 

black, matt, 0- 1-0-2 mm diam, or to 0-3 mm diam when tuberculate. Hymenium 35-45 pm tall; 

upper part (epithecium) dark green, K- or green intensifying, HNO3+ purple-red; remaining 

(lower) part ± hyaline or dilute greenish with dark green, vertical streaks, sometimes with a few, 

minute, purple-violet (K-l- aeruginose) granules. Asci clavate, 35-40x10-14 pm. Spores ovoid 

or oblong-ovoid, upper cell broader than the lower and with a more rounded apex, 1-septate, or 

rarely with a thin, additional septum in the lower cell, 7-13(-14)x(2-3-)3-4-5 pm. Paraphyses 

rather scanty, of two types: p. max. p. branched and anastomosing, haline, thin, 0-6-0-8 pm 

wide, apices often with thickened, pigmented walls and then to 2 pm wide; /?. min. p. scattered 

or in small fascicles, simple, with thickened, pigmented walls throughout and 1-5-2 pm wide, 

apices sometimes widening to 3 pm. Many apices overtopping the asci to form a ± distinct 

epithecium. Hypothecium 20-90 pm tall, dark green or dark purple (colours often intermixed), 

K-l- dark olive-green or aeruginose, HNO3-I- purple-red; hyphae interwoven, hyaline or 

thickened with pigment, 1-2 pm wide; ascogenous hyphae with swollen cells to 4 pm wide. 

Excipulum indistinct, sometimes evident as a narrow, dark greenish zone, c. 5-12 /am wide, 

forming a lateral border to the reflexed edge of the hymenium. 

Pycnidia usually present but inconspicuous, immersed between the wood fibres or emergent 

to sessile, black, with dark greenish walls, K - or green intensifying, HNO3-I- purple-red; of 

two types (a) c. 40 pm diam; conidia (mesoconidia) cylindrical, often faintly biguttulate 

3-8^-7x 1-3-1-8 pm; (b) c. 20-40 pm diam; conidia (microconidia) narrowly cylindrical, 4-5x 

0-8-1 pm. 

Chemistry: Apothecia sections C— ; material insufficient for analysis by t.l.c. 

Observations: Micarea contexta is characterised by its endoxylic thallus, very small, black, ± 

globose apothecia, dark green epithecium, dark greenish or purplish hypothecium, ovoid, 

1-septate spores, thin hyaline paraphyses and an absence of stalked pycnidia. It is apt to be 

confused with diminutive, immature forms of M. melaena, but that species has more numerous 

paraphyses, longer microconidia, and a superficial, granular thallus which contains gyrophoric 

acid when in a healthy condition. When mature, M. melaena has more robust apothecia and 

larger, 3-septate spores. M. eximia can be distinguished from M. contexta by its more brightly 

coloured epithecium and narrower, ± fusiform spores; and M. nigella can be distinguished by its 

simple spores and stalked pycnidia. M. olivacea differs in having more numerous paraphyses 

which are broader (1-1-2 îm) when hyaline, a complete absence of purple pigmentation in its 

apothecia, and relatively narrower, ± oblong spores. 

1


Habitat and distribution: M. contexta occurs on conifer lignum, probably in sheltered, 

woodland situations. Species associated with M. contexta on the specimens examined include 

Arthonia helvola, Cetraria pinastri, Cladonia spp., (scattered squamules), Hypogymnia phy- 

sodes, Lecanora symmicta agg., Lecidea efflorescens, L. pullata, Micarea anterior, M. misella, 

Parmeliopsis ambigua, and P. hyperopta. 

The known distribution of M. contexta is restricted to middle Sweden, from where it was 

found in several localities by Hedlund. Supposed collections from outside Sweden seen by me 

belong to M. melaena. Nevertheless, M. contexta should be sought for in other regions, 

especially those containing naturally occurring coniferous forests. 

Exsiccata: Malme Lich. Suec. 28 (M, S). 

9. Micarea crassipes (Th. Fr.) Coppins, comb. nov. 

(Figs4B, 12B,41C) 

Helocarpon crassipes Th. Fr., Lich. arctoi: 178 (1860); and in Nova Acta R. Soc. Sclent. Upsal. Ill, 3: 278 

(1861). - Lecidea crassipes (Th. Fr.) Nyl. in Flora, Jena 45: 464 (1861). Type: Norway, Finnmark, 

Aldjok, 1857, Th. M. Fries (S-lectotype!; isolectotypes: BM!, BM ex K!, LD, UPS!). 

Lecidea crassipes f. morlformls Th. Fr., Lich. Scand. 2: 520 (1874). Type: Norway, Finnmark, Masoy, 16 

vii 1864, Th. M. Fries (UPS-lectotype!). 

Lecidea crassipes f. pulverula Th. Fr., Lich Scand. 2: 520 (1874). Type: Norway, Sor-Trondelag, Oppdal 

hd., Dovre, Kongsvoll, Hogsnyta, 11 viii 1863, Th. M. Fries (UPS-lectotype!). 

Lecidea hypopodla f. subasslmllata Nyl. in Bull. Soc. linn. Normandle, IV, 1: 242 (1887). Type: U.S.A., 

Alaska, Bering Strait, St Lawrence Island, 1879, E. Almqulst (H-NYL 20915 - holotype!). 

Thallus growing on bryophytes or plant debris, rarely spreading on to sandy soil, composed of 

small, ± globose areolae. Areolae cream-white, pale grey-brown, sometimes ash-grey in part, 

matt, rather fragile, 0-07-0-2 mm diam; the larger areolae sometimes budding to produce 

smaller, secondary granules thereby giving the thallus an isidiose appearance. Areolae in section 

without a hyaline covering layer; thallus hyphae entirely hyaline, c. 2-3 pm wide. Phycobiont 

micareoid, cells 4-7 pm diam. 

Apothecia numerous, sessile but (in section) markedly constricted below, or turbinate, or 

short-stalked, 0-2-0-6 mm diam; disc matt, at first plane with a thin, often slightly raised, 

concolorous or slightly paler, often ± glossy, margin, 0-02-0-05 mm wide; disc later expanding, 

becoming convex and excluding the margin. On rare occasions apothecia become tuberculate 

and up to 1-4 pm diam. Hymenium 45-50 pm tall; upper part (epithecium) dark green or 

olivaceous and K-h green-intensifying, or purple-brown and K+ purple-intensifying; remaining 

(lower) part dilute purplish, K+ purple-intensifying or K+ sordid-green. Asci clavate, 40- 

48x11-14 pm. Spores 0(-l)-septate, fusiform-ellipsoid, (9-)10-17(-21)x(2-5-)3-4-5 pm. 

Paraphyses numerous, simple, or sparingly branched above, sometimes anastomosing, 1-1-5 

ju-m wide, sometimes widening above to 2-5 ^tm; apical walls hyaline but often surrounded by 

dense pigment. Hypothecium dark, variably pigmented: upper part (c. 30-60 pm) dark 

purple-brown, K-l- dark olive-green, or K-l- purple-intensifying in part or in whole; lower part 

(including 'stipe') paler and mottled, brown- or vinose-red and K- , or with purple tinge and K+ 

purple-intensifying; hyphae in upper part tightly interwoven but becoming vertically orientated 

towards the hymenium, c. 1-5-2 pm wide, embedded in a densely pigmented gel-matrix, 

intermixed with ascogenous hyphae c. 2-5-4 ^tm wide; hyphae in lower part loosely interwoven, 

becoming outwardly directed laterally (i.e. as they approach the excipulum), c. 1-1-5 pm wide, 

embedded in a gel-matrix that swells and partially dissolves in K. Excipulum well developed, 

and conspicuous in young mature apothecia, c. 50 pm wide laterally; innerpart concolorous with 

upper hypothecium; outer part dilute to dark olivaceous, K+ green-intensifying; hyphae 

radiating, branched and anastomosing, c. 1-1-5 pm wide. Excipular and hypothecial tissues 

usually extended vertically downwards to form a 'stipe' (Fig. 4B). 

Pycnidia rare, sessile, black, c. 60-150 pm diam, ostiole sometimes gaping; walls c. 10- 

17 pm wide, green-black (K- or K+ green-intensifying) in upper parts, changing to dark 

red-brown (K-) below; conidiogenous cells elongate-ampulliform, strongly tapered towards


the neck, 6-9x3-4 /xm, neck 1-1-3 /xm wide; conidia {7 mesoconidia) cylindrical, 4-5-5-7xl-2- 

1-5 ixm. 

Chemistry: Thallus K— , C-, KC-, PD-; t.l.c: no substances. 

Observations: The combination of a finely granular (sometimes isidiose) whitish to grey, 

muscicolous thallus and turbinate or short-stalked, marginate (when young), black apothecia 

make this an easy species to recognise, even with a hand lens. Internally, the apothecia have a 

well differentiated excipulum and a distinctly two-zoned hypothecium. This structure, which is 

rather 'advanced' for a Micarea, has led me to deliberate on the merits of the monotypic genus 

Helocarpon Th. Fr. However, characters such as the basic arrangement, organisation and form 

of apothecial hyphae (including paraphyses), ascus structure, morphology of pycnidia, and 

thallus structure (including phycobiont), are still those of a Micarea. The vertical extension of 

excipular and hypothecial tissues to produce a 'stipe' is not unique to M. crassipes, and is found 

in some specimens of M. lignaria and M. ternaria, and in an undescribed species from New 

Zealand (North Island: Trounson Kauri Reserve, on hepatics on rotten stumps 1967, D. M. 

Henderson (E)). This last species closely resembles M. crassipes (e.g. apothecia marginate when 

young, apothecial pigmentation, thallus type) but has broader, ellipsoid or ovoid, 1-septate 

spores (13-16-5x5-5-7 /Am) and rather sparse paraphyses. 

Overall considerations of morphology and ecology lead me to assign M. crassipes to the group 

including M. assimilata, and it is to the account of that species the reader is referred for further 

discussions. 

Habitat and distribution: M. crassipes grows over bryophytes and plant debris on the ground 

and amongst rocks in northern Fennoscandia and at high altitudes (probably above 1000 m) in 

the Alps and other central European mountains. From Norway and Sweden I have seen several 

collections from north of c. 63°N and in Russia it occurs on the Kola Peninsula. In Finland it 

extends to the southernmost parts (Karelia australis, c. 60°30'N). In central Europe it occurs in 

the western Sudety (Krkonose, at 1000 m) of Czechoslavakia, and southwards its range extends 

east to the Alps (Austrian Tirol, and Dolomiti in north Italy) and west to the Transylvanian Alps 

(Munjii Retezat, at c. 2000-2500 m) of Romania. From outside Europe it has been collected on 

St Lawrence Island in the Bering Strait; it is quite likely to have a circumpolar distribution. 

Exsiccata: Arnold Lich. Exs. 556 [A] (BM ex K, H, H-NYL 16578), 556B (BM ex K, H, H-NYL p.m. 

5105), 1121 (BM ex K). Fellman Lich. Arct. 165 (BM ex K, H, H-NYL 16570). Malme Lich. Suec. 362 

(BM). NorrlinandNyl. Herb. Lich. Fenn. 194A, B(H). Vezda L/c/i. Bohem. 282(LD). Vezda L/c/i. Sel. 11 

(BM). 

10. Micarea curvata Coppins, sp.nov. 

(Fig. 12C) 

Thallus probabiliter albido-griseus vel brunneo-griseus, granuloso-verruculosus. Algae cellulis 4-7 /Am 

diam. Apothecia immarginata, convexo-hemisphaerica mox tuberculata, atrobrunnea, 0-2-0-5 mm diam, 

aut ad 0-65 diam ubi tuberculata. Hymenium c. 60 /tm altum, ± hyalinum, cum vittis verticalibus, pallide 

fuscis; parte summa (epithecio) pallide fuscis, K-, Ascosporae fabiformes vel valde curvatae, 

(O-)l-septatae, 9-ll-7x2-5-3-8 /^m. Paraphyses ramosae et anastomosantes, graciles, c. 0-8-1 /xm latae, 

apicibus baud incrassatis et baud pigmentiferis. Hypothecium pallidum. Excipulum reflexum, pallidum, 

margine externo pallide fusescenti. Pycnidia ignota. Thallus et apothecia in sectione C+ rubra. 

Typus: Germania, Guestphalia, Gravenhorst, in muro lapideo ad lapidem arenarium, leg. Th. R. J. 

Nitschke (WRSL -holotype ) 

! 

. 

Thallus probably grey or brownish grey and granular-verrucose, but difficult to interpret in 

the single specimen in which the thallus is invaded by foreign algae and dematiaceous hyphae. 

Phycobiont micareoid, cells 4-7 pm diam. 

Apothecia convex-hemispherical to tuberculate, immarginate, dark brown, 0-2-0-5 mm diam, 

or to 0-65 mm diam when tuberculate. Hymenium c. 60 pm tall ± hyaline with pale fuscous 

brown vertical streaks; upper part (epithecium) pale fuscous brown, K- , HNO3. Asci clavate, c. 

45-50x10-13 pm. Spores fabiform or distinctly curved, (0-) 1-septate, 9-11-7x2-5-3-8 pm.


Paraphyses branched and anastomosing c. 0-8-1 /xm wide, not swollen or pigmented at apices. 

Hypothecium pale, tinged dilute straw-brown. Excipulum soon reflexed, internally pale with 

straw-brown tinge, becoming slightly darker (pale fuscous) towards the outer edge; hyphae 

radiating, branched and anastomosing, c. 0-8-1 /xm wide. 

Pycnidia not found. 

Chemistry: Sections of thallus C-l- red, PD-; sections of apothecia C-l- red; probably 

containing gyrophoric acid but material insufficient for analysis by t.l.c. 

Observations: M. curvata is characterised by its distinctly curved, 1-septate spores, fuscous 

brown pigment in the apothecia which is unchanged by K or HNO3, and C+ red reactions 

(? gyrophoric acid) of apothecia and thallus in sections. It is reminiscent of M. subnigrata but 

that species has uncurved, ellipsoid spores and all parts C-. In outward appearance M. curvata 

is similar to some forms of the almost ubiquitous Scoliciosporum umbrinum, with which it could 

be mistaken in the field. 

Habitat and distribution: M. curvata is known only from a single collection on sandstone in the 

Wroclaw Herbarium, which is labelled '?Biatora nov. sp. Auf einer Steinmauer bei 

Gravenhorst/Westfalen leg. Nitschke' (label kindly transliterated by Prof. Hertel). No recognisable 

associate species are present on the small specimen although there are a few, rounded, 

white areolae (C+ red) probably belonging to Trapelia coarctatas. ampl. From the appearance 

of the specimen it occurred in a humid, sheltered, possibly shaded or north-facing situation. 

11. Micarea denigrata (Fr.) Hedl. 

(Figs IB, 13, 42; Map 7) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 89 (1892). - Biatora denigrata Fr. in K. svenska 

VetenskAkad. Handl. 1822: 265 (1822). - Catillaria denigrata (Fr.) Boistel, Nouv. Fl. Lich. 2: 199 (1903). 

Type: Sweden, Smaland, 'Femsjo in parietibus vetustus', E. M. Fries (UPS - lectotype! [t.l.c: 

gyrophoric acid]). 

Lecidea anomala f. pyrenothizans Nyl., Lich. Scand.: 203 (1861). - Micarea denigrata f. pyrenothizans 

(Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 89 (1892). Type: Finland, Nylandia, 

Helsingfors [Helsinki], on lignum, 1860, W. A^>'/flnrfer(H-NYL 21665 -lectotype!). 

Lecidea parissima Nyl. in Crombie mJ. Bot. , Lond. 9: 178 (1871), and in/. Linn. Soc. Bot. 11: 484 (1871). 

Type: England, Middlesex, Hendon, on old pales, 1870, J. M. Crombie (H-NYL 21659 - holotype!; 

isotypes: BM!, BM ex K!). 

Lecidea spodiza Nyl. in Flora, Jena SI: 9 (1874). Type: Scotland, Perthshire, near Killin, on old worked 

conifer lignum, 1873,7. M. Crom6/e (H-NYL 21734- lectotype!; BM-isolectotypes!). 

Lecidea hemipoliella Nyl. in Flora, Jena 58: 11 (1875). - Micarea hemipoliella (Nyl.) Vezda in Vezda & V. 

Wirth in Folia geobot. phytotax., Praha 11: 100 (1976). - Micarea denigrata f. hemipoliella (Nyl.) Hedl. 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3) 78, 89 (1892). Type: Finland, Tavastia australis, Evo, 

± smooth bark oiAlnus, 1873, /. P. Norrlin (H - lectotype!, isolectotypes: H(x3)!, H-NYL 21691!). 

Lecidea discretula Nyl. in Flora, Jena 58: 444 (1875). Type: Romania, 'Circa Thermas Herculis in Banatu 

(Hung.), supra Daedaleam in trunco quercino', 1874, H. Lojka (H-NYL 21693 - holotype!). 

Lecidea aniptiza Stirton in Rep. Trans. Glasgow Fid Nat. 4: 85 (1876). Type: Scotland, Perthshire, 

Killiecrankie, on lignum, 1875, /. Stirton (GLAM - lectotype!; BM - isolectotype!). 

Lecidea denigrata var. submisella Nyl. in Vainio in Medd. Soc. Fauna Fl. fenn. 3: 112 (1878). - Catillaria 

synothea f. submisella (Nyl.) Blomb. & Forss., Enum. PI. Scand.: 92 (1880). Type: Finland, Tavastia 

australis, Korpilahti, on lignum, 1874, E. A. Lang [Vainio] (H-NYL 21661 - lectotype!). 

Lecidea spodiza f. ecrustacea Lamy in Bull. Soc. bot. Fr. 25: 440 (1878). Type: France, Puy-de-D6me, 

Monte Dore, near Cascade du Queureilh, on lignum of conifer trunk, 31 vii 1878, Lamy (H-NYL 21731 - 

lectotype!). 

Lecidea praeviridans Nyl. , Suppl. Lich. env. Paris: 5 (1879). - Biatorina praeviridans (Nyl.) Boistel, Nouv. 

Fl. Lich. 2: 194 (1903). - Catillaria praeviridans (Nyl.) Zahlbr., Cat. Lich. univ. 4: 64 (1926). Type: 

France, Haute Loire, Saugues, on Pinus bark, 18 -, A. Boistel (H-NYL 19221 - lectotype! [t.l.c: 

gyrophoric acid]; BM - isolectotype!). 

Lecidea denigrata f. sublivescens Nyl. in Flora, Jena 64: 539 (1881). Type: Romania, Maramures, 'prope


balneum kabola Pojana, com. Marmaros in Hung.', on bark of Pinus sytvestris, H. Lojka 4455, 'Lich. 

Hung. exs. (ined.) n. 305. ad int.' (H-NYL 11663 -holotype!). 

Lecidea denigrata f. pseudoglomerella Harm, in Bull. Seanc. Soc. Sci. Nancy. II, 33: 58 (1899 ['1898']). - 

Catillaria denigrata var. pseudoglomerella (Harm.) Boistel, Nouv. Fl. Lich. 2: 199 (1903). Type: France, 

Meurthe-et-Moselle, La Malgrange, on oak posts [lignum], 8 v 1894, /. Harmand, Lich, Loth. 838p.p. 

(ANGUC-lectotype! [t.l.c: gyrophoricacid]; DUKE-isolectotype!). 

Catillaria prasina f. /o«g/or Erichsen in Schr. naturw. Ver. Schleswig-Holst. 11: 101 (1937). Type: West 

Germany, Schleswig-Holstein, Lauenberg, Sachsenwald, Rev. Kl. Viert, on roots of old Fagus, 2 xii 

1934, C. F. E. Erichsen (HBO -holotype!). 

Micarea denigrata var. friesiana Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 89 (1892); 

nam. inval. (Art. 26). 

Micarea denigrata war . friesiana f. vulgaris Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 79, 90 

(1892); nom. inval. (Art. 26). 

Micarea andesitica Vezda in Poelt & Vezda, Bestimmungsschl. europ. Flechten. Erganzungsheft I: 160 

(1977); nom. nudum (Art. 32). Spec, orig.: Hungary, Matra, Matrafiired, Pipis-Legy, 380 m, on 

andesitic rock, 17 xi 1976, A. Kiszely & A. Vezda (hb Vezda!). 

? Lecidea hemipoliella* semialbula Stirton in Rep. Trans. Glasgow Fid Nats. 4: 89 (1876). - Biatorina 

synothea var. semialbula (Stirton) A. L. Sm., Monogr. Br. Lich. 2: 122 (1911). Type: Scotland, 

Sutherland, Altnaharra, on hgnum, Stirton (not seen; not traced in BM or GLAM). 

?Catillaria synothea f. major B. de Lesd., Rech. Lich. Dunkerque, Suppl. 1: 118 (1914). Type: France, 

Nord, Dunkerque, Ghyvelde, on piece of wood (Pinus) on the sand in a dune, B. de Lesdain (not seen). 

?Catillaria synothea f. fusca B. de Lesd., Rech. Lich. Dunkerque, Suppl. 1: 118 (1914). Type: France, 

Nord, Dunkerque, St-Pol, on a stake, B. de Lesdain (not seen). 

Lecidea synothea auct., non Ach. (1808). See note below. 

Note: Lecidea synothea Ach. in K. svenska VetenskAkad. Handl. 1808: 236 (1808). From the comments 

on Acharian specimens by Hedlund (1892: 91) and my examination of Acharian material in BM, it is clear 

that this name was based on material of the common and widely distributed species generally known as 

Bacidia umbrina (Ach.) Bausch or Scoliciosporum umbrinum (Ach.) Arnold (basionym: Lecidea 

umbrina Ach, Lich. Univ. : 183 (1819)). In the 19th century 'synothea' was commonly applied to Micarea 

denigrata. Despite Hedlund's comments, this misapplication has been continued by some authors, even in 

recent years (e.g. Ozenda & Clauzade, 1970: 402). To avoid the confusion that would arise if 'umbrina" 

were replaced by 'synothea' , a formal proposal to reject the name Lecidea synothea Ach. (and combinations 

from that name), under the provision of Art. 69.1, will be presented elsewhere. 

Thallus effuse and often widespreading, sometimes endoxylic but usually developing on the 

surface of the substratum (especially in the vicinity of apothecia) as convex to irregularly 

subglobose areolae. Areolae 60-200 ixm diam, greenish white to greenish grey, matt; in section 

without an amorphous covering layer, external hyphae hyaline but surrounding gel matrix often 

with dilute olivaceous, K-l- violet pigment. Thallus sometimes scurfy and dark grey-brown to 

blackish, owing to invasion by dematiaceous fungi and non-lichenized algae. Phycobiont 

micareoid, cells c. 4—7 /u,m diam. 

Apothecia usually present and numerous (see 'observations' below), scattered to confluent, 

broadly convex to subglobose, sometimes tuberculate, 0- 15-0-5 mm diam, or to 0-6 mm when 

tuberculate; immarginate, or sometimes young apothecia with an indistinct, shallow marginal 

rim, paler than the disc; disc pallid to brown or piebald (shade forms), more usually dark grey or 

black, matt. Hymenium (25-)30-40 /o-m tall, dilute olivaceous or dull brownish, K-l- violet; 

pigment often concentrated in upper part (with lower part ± hyaline), and confined to the 

gel-matrix. Asci clavate, 28-36x9-12 jxm. Spores oblong-ellisoid, oblong-ovoid, fusiform or 

bacilliform, often slightly curved, (0-)l -septate, upper cell usually slightly shorter and broader 

than the lower, (7-)9-16(-18)x2-3-3(-3-5) /u,m. Paraphyses numerous, branched and sometimes 

anastomosing, l-l-5(-l-7) /i,m wide; apices scarcely wider, and never with closely 

adhering pigment. Hypothecium 60-110 /xm tall, hyaline or very dilute yellowish straw; hyphae 

c. 1-1-5 /am wide, interwoven or some vertically orientated in upper part, intermixed with 

short-celled, ascogenous hyphae hyphae c. 2-4 /xm wide. Excipulum indistinct, but usually 

evident in sections of young, shallow-convex apothecia, hyaline; hyphae radiating, branched 

and anastomosing, hyaline, c. 1-1-5 /xm wide. 

Pycnidia usually present and numerous, immersed in the thallus or substratum (endoxylic


forms), sometimes emergent, hyaline grey or black; walls hyaline but usually dilute olivaceous 

or brownish, and K+ violet in the upper (exposed) parts; in endoxylic forms, walls olivaceous 

(K+ violet) throughout; of three types: (a) 60-150 îm diam, ostioles eventually gaping; conidia 

(macroconidia) , curved or hamate, (l-)3-septate, 12-24xc. 1 /xm; (b) 80-160 îm diam, ostioles 

often gaping: conidia (mesoconidia) short cylindric or obovate, sometimes faintly biguttulate, 

2-8^-5(-5)xl-2-l-8 fxm often extruding from the ostioles as a conspicuous white blobs; (c) 

30-50 /xm, ostioles not gaping; conidia (microconidia) narrowly fusiform or bacilliform, 

(4-5)5-7-5x0-7-0-8/xm. 

Chemistry: Thallus K— , PD— ; sections of thallus and apothecia C+ orange-red, rarely C— 

(also, parts with dull olivaceous pigment, C+ violet). If thallus is heavily parasitised and scurfy 

the C+ orange-red reaction may be difficult to obtain and gyrophoric acid may not be detectable 

by t. I.e. 

Observations: Micarea denigrata is a common and extremely polymorphic species exhibiting a 

wide range of genotypic and phenotypic variation. Its thallus can vary from being completely 

endoxylic to forming a thick, granular-verrucose, areolate crust. The colour of the apothecia 

varies from pallid, through grey or brown, to black in a ± direct relationship with exposure to 

Ught. Specimens may be abundantly fertile with numerous, large, mature apothecia. On the 

other hand, they may be ± sterile with just a few, scattered immature apothecia (or even none at 

all) but with numerous pycnidia containing mesoconidia; alternatively the latter may be 

replaced by pycnidia containing microconidia and, or, macroconidia. Indeed, all possible 

combinations of anamorphic states have been found and a few collections (e.g. Coppins 1888 

and 8384) have all three states, plus apothecia, on the same thallus. Spore length is usually in the 

range of 9-14 /^tm, but extreme forms are known in which the range is 8-10 /x,m, or 12-18 ^m. 

Gyrophoric acid is usually present in the thallus and/or the apothecia (and easily demonstrated 

by tests with C), but in some specimens its concentration is low and only detectable with 

certainty by t.l.c. ; in a few diminutive, endoxyhc forms I have been unable to detect gyrophoric 

either by a C test, or by t.l.c. In short, M. denigrata exhibits extreme variation. Despite this, with 

care and patience the species is rarely difficult to separate from other, similar species. The 

biggest problems I have experienced are where the decision has to be made between what could 

be a diminutive form of M. denigrata with small spores, or a form oiM. misella with an unusually 

high proportion of 1 -septate spores and without its characteristic stalked pycnidia (that contain 

mesoconidia). Such decisions are settled by the careful observation and measurement of 

paraphyses, and microconidia (see couplet 11 of the main key and Table 7). Shade forms of M. 

denigrata with brown, ± globose apothecia have been confused with M. elachista, but the latter 

has a brown (K+ dissolving) epithecial pigment, a more complex thallus structure (with a 

'cortex' and amorphous covering layer), distinctive pycnidia, and always lacks gyrophoric acid 

(sections of thallus and apothecia C— ). Forms of M. denigrata with a scurfy-granular thallus can 

be similar to M. prasina, but microscopic examination will show the thallus appearance to be 

caused by its disruption by invading foreign fungi and algae, and not by the presence of the small 

± discrete granules (goniocysts) characteristic of M. prasina. Furthermore, M. prasina never 

gives C+ orange-red (gyrophoric acid) reactions, and it usually has broader, rarely curved 

spores with more rounded apices. The closest relative to M. denigrata is M. nitschkeana, which is 

± identical with regard to thallus structure, pycnidial types, apothecia structure, pigmentation, 

and chemistry. Morphologically the only important difference is that the mature spores of M. 

nitschkeana are 3-septate and more consistently curved (cf. Figs 13 and 24B). In addition, the 

two species differ in their preferance of substrata: M. denigrata favouring lignum or dead bark of 

old tree trunks or stumps, and M. nitschkeana favouring corticate twigs and small branches of 

trees and shrubs. However, there is some degree of overlap as M. nitschkeana is occasionally 

found on the lignum of fence posts, but, on the other hand, I have never encountered M. 

denigrata on attached living, corticate twigs. In the field, well developed specimens of M. 

denigrata sometimes have a superficial resemblance to M. cinerea, M. lignaria, and M. 

peliocarpa, but these three species have K— hymenia and larger, 3 (or more)-septate spores. 

Diminutive endoxyhc forms of M. denigrata are indistinguishable in the field from the several,


predominantly lignicolous species with small, black apothecia, but all such species (except M. 

misella as discussed above) appear very different when examined microscopically. 

Habitat and distribution: M. denigrata is commonly found on the lignum of fallen trunks and 

old stumps of broad-leaved and coniferous trees, especially in rather open situations at the edges 

of woodlands, woodland glades, and hedgerows, etc. The communities in which it occurs are 

difficult to define, but to give some indication the following list of associated species has been 

made from collections from fallen decorticate trunks of Pinus in the native pine-woods of 

Scotland: Buellia griseovirens, Cladonia coniocraea, C. macilenta, Hypogymnia physodes, 

Lecanora expallens, L. symmicta agg., Lecidea aeruginosa, L. icmalea, Micarea peliocarpa, 

Mycoblastus sterilis, Parmelia saxatilis, Platismatia glauca, Pseudevernia furfuracea, Xylographa 

abietina, and X. vitiligo. 

M. denigrata is a very successful primary coloniser of untreated timber-work. It also occurs on 

the same substrate whose preservative has lost its effectiveness, or whose painted surface has 

flaked off. On such substrata M. denigrata has been found in a wide range of situations, e.g. 

gates, fence posts and rails, garden furniture, picnic tables and seats, window frames of 

greenhouses, wooden roof tiles (shingles), telegraph poles, and the woodwork of old carts and 

farm machinery. When on such worked wood it often forms almost pure stands, and is often 

present as a form with few apothecia but numerous pycnidia containing mesoconidia; the 

conidia often extruding as white blobs easily visible to the unaided eye or through a x 10 lens. 

Lichens associated with M. denigrata on worked wood in the British Isles include Buellia 

punctata, Cyphelium inquinans, Hypogymnia physodes, H. tubulosa, Lecanora conizaeoides, L. 

Map 7 Micarea denigrata # 1950 onwards O Before 1950


dispersa agg., L. piniperda, L. pulicaris, L. saligna, L. symmicta agg., L. varia, Lecidea 

aeruginosa, L. icmalea, Mycoblastus sterilis, Parmelia sulcata, Scoliciosporum chlorococcum, S. 

umbrinum, Strangospora moriformis, and Thelocarpon laureri. 

Further evidence of the pioneering abilities of M. denigrata comes from the find (Coppins 

1888) of it on hardboard lying in a dune-slack, and the observations by Poelt (1977; as M. 

hemipoliella) on its colonisation of dead leaves of Cladium mariscus in a fen in Bavaria. Other 

reports on soft vegetable matter are wanting, although I have found it on decaying mosses (with 

Cladonia chlorophaea agg.) on the slope of a stable sand dune on Holy Island in Northumberland 

{Coppins 4456). 

M. denigrata is occasionally found on the rather dry and loose bark of old trees (e.g. Acer 

pseudoplatanus, Alnus, Betula, Castanea, Sambucus, tjlmus, and PmM-s), usually towards their 

bases; however occurrences on ± smooth bark are very rare (e.g. type material of Lecidea 

hemipoliella, on Alnus in southern Finland). To my knowledge M. denigrata never occurs on the 

healthy, living twigs of trees, or shrubs (see 'observations' above). There are two British 

collections on shaded sandstone, one from Fife in a conifer plantation, the other from Yorkshire 

in a ditch embankment; associated species included Lecidea granulosa agg. , L. icmalea, Micarea 

peliocarpa, and Parmeliopsis ambigua. In addition, it has been found growing with Psilolechia 

lucida on the east-facing vertical side of a tomb in a Suffolk churchyard. 

M. denigrata is widely distributed in mainland Britain, but has not yet been reported from 

Ireland. In mainland Europe it is again widespread although I have not seen material from north 

of about 67°N; its range extends into eastern Europe and the Balkans and continues to the 

Caucasus. I have examined North American material from Newfoundland, Washington (state), 

and Colorado, thus suggesting a widespread distribution in that subcontinent. 

Exsiccata: Anzi Lich. Ital. 256 (BM). Arnold Lich. Mon. 46 (BM ex K). Britz. Lich. Exs. : 464 

p.p. (H). 

Fries Lich. Suec. 98 (E, UPS). Harm. Lich. Loth. 838 (ANGUC, DUKE). Hepp Flecht. Eur. 14 (E). 

Johnson Lich. Herb. 373 (BM). Krypt. Exs. Vindob. 3153 (BM, BM ex K), 3651 (BM, BM ex K, M), 4858 

(GZU). Kutak Lich. Bohem. 205 (O), 516 (O), 517 (O). Malbr. Lich. Norm. 387 (M). Malme Lich. Suec. 

145 (M, S). Migula Crypt. Germ. 132 (BM, E, MANCH). Mougeot & "HcsiXcx Stirpes Crypt. 1329p. min. p. 

(BM). Norrlin &. Nyl. Herb. Lich. Fenn. Ill (BM, H), 745 (BM, H). Poelt Lich. Alp. 22 (BM, M, WIS). 

Rabenh. Lich. Eur. 626 (BM, BM ex K, E). Samp. Lich. Port. 132 (LD). Vezda Lich. Sel. 1430 (BM). 

Weber Lich. Exs. 73 (E, DUKE, M, NMW, WIS). Zwackh Lich. Exs. 394 (BM ex K, H). 

12. Micarea elachista (Korber) Coppins & R. Sant. , comb. nov. 

(Figs IC, 14A) 

Biatora elachista Korber, Parerga lich.: 159 (1860). - Catillaria elachista (Korber) Vainio in Acta Soc. 

Fauna Fl. fenn. 57 (2): 455 (1934). Type: Germany, Baden-Wiirttemberg, Heidelberg, on old trunk of 

Castanea sativa, W. E. von Ahles (L 910, 138-100-lectotype!; L 910, 138-32-isolectotype!) See note 

below. 

Lecidea anomala *L. glomerella Nyl., Lich. Scand.: 203 (1861). - Biatorina glomerella (Nyl.) Arnold in 

Flora, Jena 53: 474 (1870). - Catillaria glomerella (Nyl.) Th. Fr., Lich. Scand. 2: 578 (1874). - Micarea 

glomerella (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 75, 85 (1892). Type: 

Finland, Ostrobottnia australis, Botom, 1859, A. J. Malmgren (H-NYL 19123 - lectotype!). 

Lecidea poliococca Nyl., Lich. Scand. : 203 (1861). - Catillaria denigrata f. poliococca Vainio in Acta Soc. 

Fauna Fl. fenn. 57 (2): 460 (1934). - Catillaria synothea var. poliococca (Nyl.) Erichsen in Annls mycol. 

41: 205 (1943). Type: Sweden, Uppland, 'Upsalia', in silva 'Parken', ad pinos decorticatos' , 1852, W. 

Nylander (H-NYL 19144- holotype!). 

Lecidea sororians Nyl. in Flora, Jena 58: 445 (1875). - Bacidia sororians (Nyl.) H. Olivier in Bull. Geogr. 

hot. 21: 168 (1911). Type: Finland, Tavastia australis, Korpilahti, near Raianlahti, on rock with 

Stigonema sp., 1873, E. A. Lang [Vainio] (H-NYL 17234 -holotype!). 

Lecidea glomerella f. simplicata Nyl. in Vainio in Medd. Soc. Fauna Fl. fenn. 10: 28 (1883). Type: Finland, 

Tavastia australis, Evo, 'supra truncum pineum', 1874, J. P. Norrlin, Norrlin & Nyl. Herb. Lich. Fenn. 

314 (H- lectotype!; isolectotypes: BM!,H!M!). 

Lecidea glomerella var. poliococcoides Vainio in Medd. Soc. Fauna Fl. fenn. 10: 29 (1883). Type: Finland, 

Karelia borealis, Lieksa, Vieki, on burnt lignum, 1875, E. A. Vainio (TUR-VAINIO 22326 - holov 

type!).


Catillaria elachista var. carbonicola Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 458 (1934). Type: Finland, 

Tavastia australis, Korpilahti, 'hiiltyneella kannolla', 1873, E. A. Lang [Vainio] (H-NYL 19143 - 

isotype!). 

Notes on the typification o/Biatora elachista Korber. In the protologue to B. elachista, Korber (loc. cit.) 

gives the following habitat and locality information: 'On alten Striinken der Castanea vesca bei Heidelberg 

von Hrn. v. Zwackh und Ahles aufgefunden.' Material borrowed from the Korber Herbarium (L) 

consisted of four specimens: i. bei Heidelberg, Zwackh, annotated: 'Biatora elachista Kbr. nov. sp.' (L 

910, 138-101). ii. Heidelberg, Ahles (L 910, 138-100). iii. identical label as ii. (L 910, 138-32). iv. 

Forstrevier Goleow bei Rybnik, iii 1872, B. Stein (L 910, 138-298). (i-iii) are on Castanea lignum and (iv) is 

on conifer lignum. 

The most obvious choice for lectotype is (i) but it is not a Micarea and does not conform to the usual 

interpretation of the name (e.g. Vainio, 1934: 45; Ozenda & Clauzade, 1970: 401). I do not know the 

identity of this species, but it may be close to Lecidea hypopta Ach. ; a brief description follows: 

Thallus probably mostly endoxylic but obscured by a pale farinose crust of a non-lichenised alga. 

Apothecia numerous, reddish brown to brown-black, epruinose 0- 15-0-3 mm diam. Young 

apothecia with a pale margin which is soon excluded as the disc expands and becomes convex. 

Epithecium reddish brown, turning olivaceous brown in K. Hymenium 30-35 /u,m tall. Paraphyses 

simple or forked in their upper part, c. 1-1-5 ^tm wide; upper 5-15 ^tm often with pigmented walls 

and up to 2-5 /x,m wide. Asci clavate 28-35x10-11 /xm, 8-spored. Spores simple or l-(rarely 2-) 

septate oblong-ellipsoid, straight or slightly curved, contents often becoming brown, 8-13x3-4 /u,m. 

Excipulum hyaline within, reddish brown at edge, often with a penetrating algal layer c. 25 )u,m wide. 

Algal cells c. 7-13 ^tm diam. Pycnidia not found. 

Specimens (ii) and (iii) are presumably those that Korber attributed to Ahles in the protologue and are 

therefore syntypes. Both belong to the species generally known as Catillaria elachista (Korber) Vainio, and 

it is with these that the name is lectotypified here. Specimen (iv) is not a syntype and does not belong 

to either of the above taxa. It has small, brown thinly white-pruinose apothecia, a dark reddish 

brown hypothecium and small, ellipsoid spores, 7-8x2-5-3 /am, and is referable to Lecidea apochroeella 

Nyl. 

Thallus effuse, superficial, consisting of dispersed to continuous, convex to subglobose 

areolae; areolae greenish white or whitish grey, sometimes tinged grey-brown or olivaceous, 

occasionally dark brown (when on burnt lignum), matt, sometimes ± white-pruinose, c. 

0-08-0-16(-0-25) mm diam. Areole in section (Fig. IC) with a c. 10-12 /u,m tall, hyaline or 

greyish (pigment in gel-matrix, K+ violet), algal-free 'cortex', composed of interwoven, hyaline 

hyphae (c. 1-5 ^tm wide) that separate in K; outer surface of cortex sometimes bound by a 

hyaline, amorphous, densely gelatinised layer ('epicortex'), c. 3-5 /xm tall. This organised 

structure with a 'cortex' and 'epicortex' is often disrupted by the invading torulose hyphae of a 

dematiaceous hyphomycete. Phycobiont micareoid, cells 4-7 jxm diam. 

Apothecia usually numerous, immarginate, convex to ± globose, often becoming tuberculate, 

dark brown to brown-black, matt, (0-08-)0- 12-0-3 mm diam, or to 0-8 mm diam when 

tuberculate. Hymenium 30-40 />tm tall. Upper part (epithecium) usually well defined, up to 

12 jxm tall, dark fuscous-brown; pigment concentrated into dense amorphous clumps, but 

dissolving and fading into solution (without changing colour) in K, HNO3— and not dissolving; 

epithecium of young apothecia sometimes Kf+ violet due to the additional presence of the dull 

olivaceous, K-l- violet pigment which occurs in greater concentrations in the pycnidia. Remain- 

ing (lower) part of hymenium, hyaline with dilute yellowish brown vertical streaks. Asci clavate, 

23-35x10-12 jxm. Spores fusiform, oblong-fusiform or ovoid-oblong, often slightly curved, 

mostly 0-1-septate and slightly constricted at the septum, (9-)ll-15(-19)x2-3'5 jxm; occasionally 

becoming 2- or 3-septate and up to 20-24 /xm long; old spores sometimes with brownish 

contents. Paraphyses numerous, hyaline throughout, 0-8-1 /u,m wide in mid-hymenium but 

gradually widening above to l-7(-2) /xm; sparingly branched and sometimes anastomosing, but 

becoming richly branched above where their entangled apices, together with the brown 

pigment, form a ± well delimited epithecium. Hypothecium 60-150 /xm tall, pale, tinged dilute 

yellowish brown, K— , HNO3-; hyphae hyaline, 1-1-5 /xm wide, interwoven, becoming ± 

vertically orientated towards the hymenium; ascogenous hyphae with swollen cells, mostly 2-


4 fim wide. Excipulum poorly developed and much reflexed, sometimes evident as a narrow, 

pale fuscous-brown zone; hyphae hyaline, radiating, branched and anastomosing, 1-1-5 /xm 

wide. 

Pycnidia usually present and numerous, developing from within areolae but soon becoming 

emergent and sometimes ± sessile; of two types: (a) c. 100-200 /xm diam, grey-brown but 

usually paler or whitish around the ostiole, surface smooth and ± glossy; ostioles distinct and c. 

20 jxm diam, sometimes gaping and to 50 jxva diam; wall not continuing below the base, laterally 

c. 23-40 /xm wide, dilute olivaceous or brownish, K-l- violet, and formed of interwoven hyaline 

hyphae (c. 1-5 /xm wide) that ± separate in K; conidiogenous cells cylindrical, occasionally 

percurrently proliferating, c. 4—7x1-2 /xm; conidia (mesoconidia) ± cylindrical, sometimes 

slightly wider at proximal end, often faintly biguttulate, 3-5-4-5xl-3-l-7(-2) /xm. (b) similar in 

appearance and structure to above, but smaller and 60-100 /xm diam; conidiogenous cells 

cylindrical, c. 3-6x1 /xm; conidia (microconidia) narrowly cylindrical, (4-)4-5-6(-6-5)x 

0-7-1 /xm. 


by t. I.e. 

, KC— , PD— ; sections of apothecia C— ; no substances detected 

Observations: The combination of ± globose areolae, ± globose, brown apothecia, dense 

brown (dissolving in K) epithecial pigment, C- hymenium, fusiform, 1-3-septate spores and 

distinctive, ± glossy pycnidia usuall-y make this species easy to identify. Some confusion has 

been made with shade forms of M. denigrata, but these can be distinguished by their rather 

adnate and often larger apothecia, C-f- orange-red (gyrophoric acid) hymenium and thallus, 

never glossy, thin-walled pycnidia, and, when present, curved macroconidia. M. elachista is 

closely related to M. rhabdogena (q-v.), which differs in having an endoxylic thallus, smaller, 

mostly simple, spores, and black pycnidia. 

The areolae of M. elachista sometimes have a white-pruinose appearance: the 'pruina' 

resulting from the partial disintegration of the thin 'epicortex'. 

Habitat and distribution: M. elachista is found, often in the company of Parmeliopsis spp. , on 

the lignum (rarely bark) of partially or wholly decorticate trunks or large stumps of old trees, 

especially Castanea, Pinus, and Quercus. Most collections from France and Germany were 

made from Castanea, whereas those made from Scandinavia were mostly from conifers. It 

occasionally occurs on burnt or charred stumps and my collection (Coppins 6017) from Sweden, 

on Pinus, was accompanied by Chaenotheca ferruginea, Hypocenomyce friesii, Lecidea granulosa 

agg. , Micarea melaena, and Parmeliopsis spp. It seems to be rare on worked timber, but was 

collected on old fence-posts in Bavaria by Arnold. 

I have seen only one saxicolous gathering of M. elachista, i.e. the holotype of Lecidea 

sororians from southern Finland, with which occurred colonies of a Stigonema sp. , a few lobes of 

a brown Parmelia (? P. verruculifera) and, according to Nylander, Biatorella torvula (not 

present in the existing material). This specimen is very small and in poor condition, and the few 

spores seen appeared to be abnormally developed, 1-3-septate, 15-24x2-2-5 /xm; other 

features of the thallus and apothecia agree with M. elachista (pycnidia were not found) and I 

think it is most unlikely to represent a distinct taxon. 

M. elachista appears to avoid the more strongly oceanic areas of Europe, and is known from 

mid-Sweden, southern Finland, France (Haute Vienne, the Massif Central and the Pyrenees), 

southern Germany (Hessen, Baden-Wiirttemberg and Bavaria), and the Austrian Tirol. Its 

presence in Britain is a possibility and it should be sought for, especially in the central Highlands 

and east Scotland, and the Welsh border counties. 

Exsiccata: Arnold Lick. Exs. 1471 (BM ex K, M). Arnold Lick. Mon. 246 (BM ex K, M). Malme Lick. 

Suec. 21 (M, S). Norrlin & Nyl. Herb. Lick. Fenn. 314 (BM, H, M); 724 (BM). Vezda Lick. Sel. 1134 

(BM). Zwackh Lich. Exs. 122 (H-NYL 18828, M).


13. Micarea eximia Hedl. 

(Figsl4B,40C) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 80, 84, 95 (1892). - Catillaria malmeana Zahlbr., 

Cat. lich. univ. 4: 56 (1926); nom. nov.; non Catillaria eximia Malme. Type: Sweden, Dalarna, Alvdal, 

Hallstugen, vi 1891, J. T. Hedlund (S - lectotype! [t.l.c: no substances]; S - isolectotype!; material 

distributed as Malme Lich. Suec. Exs. 26 is possibly part of this collection). 

Thallus effuse, endoxylic, of minute clusters (c. 15-40 yun diam) of the phycobiont amongst 

the wood fibres, with intertwining hyphae that often have green walls, K-, HNO3+ red. 

Phycobiont micareoid, cells 4-7 /x,m diam. 

Apothecia numerous, immarginate, ± convex-globose from the start, often becoming 

tuberculate, black, matt or slightly glossy, 0- 1-0-2 mm diam, or to 0-4 mm diam when 

tuberculate. Hymenium 30-45 /am tall; upper part bright to dark aeruginose, K— or dulling, 

HNO3+ purple-red; remaining (lower) part ± hyaline with aeruginose vertical streaks. Asci 

clavate, c. 30-40x11-12 ^.m. Spores oblong-fusiform, sometimes slightly curved, 9-14 

(-16) X 1 •8-2-5 /xm. Paraphyses rather scanty, branched and anastomosing, thin and 0-7-0-8 /xm 

wide, but upper c. 5-17 (xm with thickened, pigmented walls and then 2-3 /xm wide; usually a few 

paraphyses have pigmented walls throughout their length and are about 2 îm wide in the middle 

hymenium. Hypothecium 45-120 //,m tall, dilute reddish- or purplish brown, sometimes with 

darker blotches, K+ dull greenish, HNO3+ purple-red, pigment confined to gel-matrix; hyphae 

interwoven, hyaline, c. 1-1-5 /xm wide; ascogenous hyphae with swollen cells up to 4 îm diam. 

Excipulum indistinct, sometimes evident as a dark green or brownish (K-l- olivaceous) reflexed 

zone with branched, radiating, pigmented hyphae c. 1-5-2 /xm wide. 

Pycnidia numerous but inconspicuous, usually immersed between surface wood fibres, but 

sometimes ± emergent, black, c. 35-80 /xm diam; walls dark green, K-, HNO3-I- purple-red, 

composed of tightly bound, pigmented hyphae, c. 2 /xm wide. Conidia (mesoconidia) cyhndri- 

cal, sometimes biguttulate, 3-9-5-5X 1-1-4 /xm. 

Chemistry: Sections of apothecia and thallus C— ; no substances detected by t.l.c. 

Observations: Micarea eximia is characterized by the combination of an endoxylic thallus, 

small, black, ± globose or tuberculate apothecia, bright green upper hymenium, reddish or 

purplish brown (K-l- green) hypothecium, simple or 1-septate, oblong-fusiform spores, and an 

absence of stalked pycnidia. It is most likely to be confused with M. contexta and M. olivacea. 

The former has broader, ovoid spores and a darker hypothecium; and the latter has a less 

brightly coloured hymenium, a darker olivaceous hypothecium, more numerous and broader 

paraphyses, shorter and slightly broader spores with rounded apices, and shorter mesoconidia. 

M. nigella differs from M. eximia in having a purplish brown upper hymenium, a darker 

hypothecium (hyphal walls pigmented), ellipsoid to oblong-ovoid, simple spores, and stalked 

pycnidia. The spores of M. eximia are similar to those of M. denigrata, but that species has an 

olivaceous, K-l- violet pigment in its upper hymenium and pycnidial walls, and a ± hyaline 

hypothecium. 

Habitat and distribution: M. eximia is a rare or overlooked species of conifer lignum, known 

only from middle Sweden and northern Finland. Associated species on the specimens examined 

include Bacidia retigena, Calicium glaucellum, Cetraria pinastri, Cladonia spp., Lecideapullata, 

Micarea misella, Parmeliopsis aleurites, P. ambigua, P. hyperopia, and Xylographa vitiligo. 

14. Micarea globulosella (Nyl.) Coppins, comb. nov. 

(Figsl5,43A-B;Map24) 

Lecidea globulosella Nyl., Lich. Jap.: 69 (1890). - Bacidia globulosella (Nyl.) Zahlbr., Cat. lich. univ. 4: 

202 (1926). Type: Japan, Yokohama, on bark, 1879, E. /4/m^Mwf(S -lectotype! ; isolectotypes: H-NYL 

17412! and 17413!, S!). 

Thallus effuse, of scattered or, more usually, ± contiguous areolae. Areolae convex, whitish 

or grey, not gelatinous when wet, 40-150 /xm diam; in section without an amorphous covering


layer, outer hyphae sometimes olivaceous (or brownish) and K+ violet. Phycobiont micareo'id, 

cells 4-7 /Ltm diam. 

Apothecia numerous, immarginate, convex and adnate to subglobose, sometimes becoming 

tuberculate, greyish or brownish black, sometimes pale grey or pallid (shade forms), 0- 1-0-3 mm 

diam. Hymenium 35-40 /xm tall, dilute olivaceous or dilute olive-brown, K+ violet. Asci 

clavate, 30-35x9-12 jxm. Spores fusiform-acicular or ± rod-shaped, slightly curved or ± 

straight, 0-3(-6)-septate, 13-26 x(l'5-)2-2-5(-3) /xm. Paraphyses numerous, branched and 

sometimes anastomosing, c. 1 fxm wide, sometimes widening to 1-5 fxm towards their apices; 

apical walls hyaline. Hypothecium 50-70 /xm tall, hyaline. Excipulum indistinct, evident in some 

sections as a narrow, reflexed, lateral border to the hymenium, hyaline or dull olivaceous (then 

K-l- violet); hyphae radiating, branched and anastomosing, c. 1 jxm wide. 

Pycnidia frequent but inconspicuous, immersed or emergent, whitish to dark grey; walls 

olivaceous or dull brownish, K-l- violet (especially around the ostioles), sometimes ± hyaline. 

Pycnidia of two types: (a) immersed within large areolae, sometimes emergent, 60-100 />im 

diam; conidia (mesoconidia) ± cylindrical, 3-6-5-3X 1-1-4 /xm; (b) similar but smaller, c. 30-40 

fxm diam; conidia (microconidia) narrowly cylindrical, 3-8-5x0-8-1 /am. 

Chemistry: Thallus C+ red, K— , PD— 

to oHvaceous pigment); t.l.c: gyrophoric acid. 

; apothecia sections C— or C-l- red (also C-l- violet due 

Observations: M. globulosella is closely related to M. denigrata and M. nitschkeana but can be 

distinguished by its longer, almost acicular, or rod-shaped spores. My early suspicions that M. 

globulosella could be a long-spored variant of M. nitschkeana were removed by the examination 

of microconidia, those of M. nitschkeana being significantly longer (mostly 5-5-7 /am). Long, 

curved macroconidia, as found in M. denigrata and M. nitschkeana, have not yet been found in 

M. globulosella, but they should be sought for in additional material. See under M. synotheoides 

for differences from that species and Bacidia beckhausii. The name Micarea bacidiella (a 

synonym of B. beckhausii) was mistakenly applied to M. globulosella by Vezda & Wirth (1976) 

and Poeh&Vezda (1977). 

Habitat and distribution: M. globulosella is a rare but widespread species, so far known to me 

from Wales, Sweden, Finland, south-east France, Bavaria (Allgau), Czechoslovakia (High 

Tatra) , Canada (Quebec) , and Japan (Honshu) . It seems to prefer conifer bark , but at the Welsh 

locality it occurred on the top of an old gate in a wooded valley, and was accompanied by 

Lecanora piniperda, Lecidea aeruginosa, and L. icmalea. The specimen from Allgauer Alpen is 

associated with Graphis scripta, Menegazzia terebrata, Stenocybe major, and Frullania sp. on 

Abies; that from Sweden is on Picea with Cetraria pinastri and Ptilidium pulcherrimum; and 

those from Czechoslovakia are on Picea, with Cetraria chlorophylla, Hypogymnia physodes, 

and Ptilidium pulcherrimum. At the French locality it was growing on an old basidiome of the 

polypore Daedaleopsis confragosa on Pinus hapelensis. 

It seems to require humid conditions in old forest situations, but has a more widespread 

distribution than M. synotheoides which is strongly oceanic. M. globulosella may have a greater 

tolerance to cold winter temperatures, such as occur in the central European montane regions. 

Exsiccata: Rasanen Lichenoth. Fenn. 426 (hb Vezda). 

15. Micarea hedlundii Coppins, sp. nov. 

(Figs 14C, 44A) 

Thallus effusus, olivaceo-viridis, subtiliter granulosus ad 0-4 mm crassus, constatus ex goniocystis; 

goniocystae c. 20-40 /u.m diam, omnes cum pigmento flavo-brunneolo, K+ purpureo-violaceo et oleoso. 

Algae cellulis 4-7 /u,m diam. Apothecia vulgo pauca vel etiam nulla, immarginata, convexo-hemisphaerica 

mox tuberculata, pallida vel griseo-fusca demum obscure fusca, 0-15-0-5 /xm diam. Hymenium c. 35 /xm 

altum, ± hyalinum vel p.p. dilute olivaceo-brunneolum, K-l- violaceum. Ascosporae ellipsoideae, 

ovoideae vel oblongae, simplices interdum 1- septatae, 6-5-10(-12)x 2-3-4 /xm. Paraphyses aliquantum 

paucae, ramosae, c. 0-7-l(-l-5) /xm latae, apicibus vix incrassatis, incoloratis, Hypothecium pallidum.


Excipulum paulum evolutum. Pycnidia numerosa, conspicua, stipitata, 0- 1-1-0 mm alta, 0-07-0-14 mm 

diam, cum stipibus simplicibus vel ramosis, toto griseo-brunneo vel roseo-brunneo cum tomento exili 

albido. Conidia oblongo-ellipsoidea vel oblongo-ovoidea, (4-)4-5-5-5(-6)xl-3-l-7 ^tm. 

Typus: Norvegia, Opplandia, par. Ringebu, ad oriento-boreo-orientum ex Ringebu, ad S0raa, inter 

Nyhamnsbekken et Ulveslabekken, alt. 400 m, ad truncum decorticatum, 25 viii 1979, leg. L. Tibell 8657 

(UPS-holotypus). 

Thallus epixylic or over bryophytes on lignum, effuse, dull olive-green, consisting of fine 

granules (goniocysts) forming a loose crust to c. 0-4 mm thick. Goniocysts c. 20-40 /xm diam, the 

centre of each with a yellow-brown or dull orange pigment which in K appears as purple-violet 

oily droplets. Phycobiont micareoid, cells 4-7 /um diam. 

Apothecia few or sometimes absent, immarginate, convex and soon becoming tuberculate, 

pallid to grey-brown or dark brown, 0-15-0-5 mm diam. Hymenium c. 35 /xm tall, ± hyaline with 

vertical streaks of straw-brown or pale olive-brown, K+ violet, C+ violet, HNO3-I- reddish; 

lower hymenium sometimes with dull orange, K+ purple-violet (oily droplets) pigment. Asci 

clavate, c. 30x12 ixm. Spores ellipsoid, ovoid or oblong, simple, a few sometimes 1-septate, 

6-5-10(-12)x2-3-4 /am. Paraphyses rather scanty, branched and sometimes anastomosing, 

0-7-l(-l -5) /Ltm wide, apices not swollen or pigmented. Hypothecium c. 70-100 /xm tall, hyaline, 

or with dull orange, K-f- purple-violet (oily droplets) pigment in upper part; hyphae interwoven, 

becoming vertically orientated towards the hymenium, c. 1-1-7 (xm wide, intermixed with 

swollen short-celled ascogenous hyphae. Excipulum indistinct and soon reflexed, sometimes 

discernible in young apothecia as a narrow, hyaline, non-amyloid zone; hyphae radiating, 

branched and anastomosing, c. 0-8-1 /zm wide. 

Pycnidia numerous and conspicuous, stalked with one or to about five borne terminally on 

simple or branched pycnidiophores. Pycnidiophores (including pycnidia) grey-brown or pinkish 

brown, covered with a thin whitish tomentum, c. 01-1 mm tall and 0-07-0-14 mm diam; lower 

parts of pycnidophores often covered in goniocysts. Pycnidiophore and pycnidial wall tissues 

composed of hyaline hyphae bound by a dilute reddish brown matrix reacting K+ violet or 

violet-brown and HNO3-I- reddish; surface of pycnidiophores and pycnidia with protruding, 

slender, flexuose, hyaline tomental hyphae c. 0-7-1 /xm wide. Conidiogenous cells ampuUiform 

to ± cylindrical, 5-lOxc. 1-5 /am, often swollen at base to 2-5 /am wide. Conidia (mesoconidia) 

oblong-ellipsoid or oblong-ovoid, (4-)4-5-5-5(-6)xl-3-l-7 /am. 

Chemistry: Thallus C— , PD— 

; apothecia sections C— (but with C+ violet pigment); no 

substances or traces of 'prasina-unknown B' (? contaminant) detected by t.l.c. 

Observations: Micarea hedlundii is readily identified by its finely granular, darkish green 

thallus with distinctly stalked, pinkish brown, tomentose pycnidia. Several other Micarea 

species have stalked pycnidia but in none of them are they tomentose. Another unique feature 

oiM. hedlundii is the dull orange pigment, present in the goniocysts and sometimes in the lower 

hymenium and upper hypothecium, which appears as purple-violet oily droplets in K. Unlike 

the olivaceous K-f violet pigment found in this species and others (e.g. M. denigrata, M. 

nitschkeana, and M. prasina), it remains unchanged in C and 50% HNO3. M. hedlundii is 

unlikely to be confused with any other Micarea, but is probably closely related to M. prasina. M. 

anterior has reddish brown, stalked pycnidia but they are glabrous and produce shorter conidia; 

in addition its apothecia and pycnidia are completely devoid of K+ violet pigments. 

The chemistry of M. hedlundii is problematical. Of the three specimens tested by t.l.c, the 

type from Norway contains no detectable substances, but the two specimens from Austria 

(GZU) and Germany (hb Poelt) appear to contain small amounts of 'prasina-unknown B'. This 

substance is known elsewhere only in the type race of M. prasina and it is possible that its 

detection in the aforesaid samples is due to contamination by that species. However, the 

production of 'prasina-unknown B' by M. hedlundii is a possibility meriting further study, 

especially as M. hedlundii and M. prasina appear to be closely related to one another. New and 

carefully collected specimens ofM hedlundii are required to resolve this problem. 

Habitat and distribution: M. hedlundii occurs on old stumps (? mainly of conifers) in 

woodlands. Few associated lichens are present on the specimens examined, mostly just a few


scattered squamules of Cladonia spp. and fragments of Lepraria spp, although one of the 

Norwegian collections includes a Chaenothecopsis. M. hedlundii seems to be a very rare species, 

but is known from scattered localities in Norway, Sweden, Germany (Bayern), Austria 

(Steiermark), and possibly Switzerland. 

Etymology: This new species is named in honour of Johan Teodor Hedlund (1861-1953) in 

recognition of his pioneering study of Micarea included in his doctorate thesis for the University 

of Uppsala (Hedlund, 1892), a work that has been a constant inspiration during my own studies. 

16. Micarea incrassata Hedl. 

(Figs4C, 16,41A-B;Map3) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 82, 94 (1892). Type: Austria, 'supra muscos in 

summo jungo Kraxentrag circa Brenner, Tirol, c. 2800 m. alt.', A. Minks (S - holotype!). 

Lecidea assimilata B. [var.] infuscata Th. Fr., Lich. Scand. 2: 522 (1874). - Lecidea assimilata f. infuscata 

(Th. Fr.) Vainio in Medd. Soc. Fauna Fl. fenn. 10: 85 (1883). Type: Norway, Sor-Trondelag, Oppdal 

hd., Dovre, Kongsvoll, Hogsnyta, 17 viii 1863, Th. M. Fries (UPS - lectotype!; UPS - isolectotype! 

[t.l.c: no substances]). 

Thallus growing on bryophytes, plant debris or sandy soil, composed of confluent, convexverrucose 

areolae that are intermixed with cephalodia. Areolae dull grey-white, or grey-brown 

to dark grey, matt, 0-08-0-3 mm diam; in section, usually without a hyaline amorphous covering 

layer (but sometimes seen in sections of young areolae), hyphae in outer c. 10 jxm often with 

light brown walls. A ± algae-free medulla often differentiated. Thallus hyphae c. T8-3 /am 

wide. Phycobiont micareoid, cells 4—7 /u,m diam. Cephalodia often present, irregularly globose 

and hidden amongst the areolae but sometimes visible externally as brown areolae-like 

structures, 0-2-0-6 mm diam; containing Nostoc, cells 3-5 /i,m diam. Less often present are 

irregular, rather loose clusters (? cephalodia) of Stigonema. 

Apothecia numerous, immarginate, convex, ± adnate and often partly immersed by surrounding 

areolae, black, matt, 0-3-0-8(-l) mm diam, sometimes forming irregular tuberculate 

clusters up to T2 mm diam. Hymenium 45-50 îm tall; upper c. 10 ixm (epithecium) dark 

aeruginose or olivaceous, K— , HNO3-I- red; remaining (lower) part dilute greenish or hyahne. 

Asci clavate, 45-48x11-14 [xm. Spores ellipsoid, oblong-ellipsoid, oblong-ovoid or oblongfusiform, 

0-l(-2)-septate, (10-)12-17x4-4-8 /xm. Paraphyses numerous, simple below, sometimes 

forked above, sometimes anastomosing, (l-)l-5-2 ixm wide, sometimes widening above 

to 3 jxm; apical walls hyaline although surrounded by dense pigment in the surrounding gel 

matrix. Hypothecium c. 150-400 /xm tall, sometimes 'rooting' to the base of the adjoining 

areolae, dark red-brown, without a purple tinge, K-, HNO3+ bright orange-brown; hyphae 

interwoven, or ± vertically arranged in upper part, c. T7-2-5(-3) ^tm wide, surrounded by 

densely pigmented matrix; ascogenous hyphae c. 2-5-5 /xm wide. Excipulum indistinct, 

sometimes evident as a reflexed, dull olivaceous or reddish brown zone; hyphae radiating, 

branched and anastomosing, c. 1-2 /xm wide. 

Pycnidia rare, immersed to sessile, black, 30-60 /am diam; walls dark green above, changing 

to reddish brown at base; conidia (? mwroconidia) cylindrical or cylindrical-fusiform, 6-9x1- 

T3/am. 

Chemistry: Thallus K-, C-, PD-; t.l.c: no substances. 

Observations: See M. assimilata. 

Habitat and distribution: M. incrassata occurs in much the same habitats as M. assimilata but 

appears to be more widely distributed. In Europe it has an arctic-alpine distribution ranging 

from Spitzbergen (c. 79°N) in the high arctic, southwards to the Kola peninsula, central Norway 

(Opland), and central Sweden (Harjedalen). Further south it occurs in the Scottish highlands 

(Clova in Angus, and near Loch Merkland in East Sutherland), Denmark, and the Alps (Upper 

Bavaria, Austrian Tirol, and Switzerland). I have not seen it from Alaska or Canada although it 

probably occurs in those regions. From further south in North America it has been collected at


altitudes of c. 3400-3900 m in the Rocky Mountain National Park in Colorado by Anderson 

(1964). M. incrassata has been collected on Kerguelen Island in the southern Indian Ocean 

(48°30'S) and is so far the only arctic or arctic-alpine Micarea known to have a bipolar 

distribution. 

Exsiccata: Fellman Lich. Arct. 164 (BM ex K, H, H-NYL 16567), 166 (BM ex K, H). 

17. Micarea intrusa (Th. Fr.) Coppins & Killias, comb. nov. 

(Figs 17, 55; Map 22) 

Lecidea intrusa Th. Fr. in Bot. Notiser 1867: 152 (1867). - Catillaria intrusa (Th. Fr.) Th. Fr., Lich. Scand. 

2: 579 (1874). - Lecidiopsis intrusa (Th. Fr.) Zopf in Hedwigia 35: 338 (1896). - Conida intrusa (Th. Fr.) 

Sacc, Syll. Fung. 18: 187 (1906). Type: Finland, Tavastia australis, Mustiala, 1867, A. Kullhem (UPS- 

holotype!). 

Lecidea aphanoides Nyl. in Flora, Jena 51: 476 (1868). Type: Scotland, South Aberdeenshire, Braemar, 

'Craig Guie', viii 1868, J. M. Crombie (H-NYL 20237 - holotype!); supposed isotype material in BM 

appears to be all Scoliciosporum umbrinum (Ach.) Arnold. 

Lecidea melaphana Nyl. in Flora, Jena 52: 83 (1869). Type: Scotland, South Aberdeenshire, Braemar, 

'Craig Guie', viii 1868, J. M. Crombie (BM - isotype!). 

Lecidea contrusa Vainio in Medd. Soc. Fauna Fl. fenn. 10: 29 (1883); nom. illeg. (Art. 63). 

Thallus occasionally effuse but more often forming small patches (to c. 1 cm diam) amongst 

other lichens, irregularly granular-verrucose and often rimose, sometimes forming discrete 

areolae (c. 0- 1-0-3 mm diam) which may coalesce to form 'composite areolae' up to c. 1 mm 

diam and 0-5 mm tall. Thallus dark olive-grey or brownish grey; surface matt, and usually scurfy 

due to invasion of free-living algae. In section, sometimes with a hyaline amorphous covering 

layer up to 12 /xm thick but this is usually disrupted by invading algae ; walls of uppermost hyphae 

sometimes thickened with olive-green pigment, K-, HNO3-I- red; phycobiont layer c. 70^100 

mm thick, above a ± distinct hyaline medulla. Phycobiont not micareoid; cells globose, large, 

7-21 /xm diam, with thick hyaline walls 1-2 /xm thick, each cell deeply penetrated by a 

haustorium of the mycobiont (Fig. 55). 

Apothecia numerous, immarginate, convex, adnate, black, or slightly glossy when young, 

0-14-0-4 mm diam. Hymenium 40-50 /xm tall; upper (c. 20 /x,m) part aeruginose-green, K-, 

HNOs-t- red; remaining (lower) part ± hyaline. Asci clavate, 35-45x14-15 /xm; cytoplasm of 

asci and spores sometimes with an orange, K+ purple-red pigment. Spores elhpsoid or 

ovoid-ellipsoid, occasionally oblong-ellipsoid or oblong fusiform and then sometimes slightly 

curved, simple or 1-septate and septum often eccentric, rarely 3-septate (7-)9-14(-17)x 

(4-)4-5-5-5(-6) /am. Paraphyses numerous, branched and anastomosing, c. l-l-5(-l-7) /xm 

wide; apices not swollen or pigmented (green pigment confined to the surrounding gel-matrix). 

Hypothecium c. 110-200 /u,m tall, hyaline or with very faint olivaceous brown tinge in places. 

Fig. 55 Micarea intrusa: a phycobiont cell penetrated by a haustorium of an associated mycobiont hypha. 

Scale = 10 /u,m.


upper part sometimes pale orange, K+ purple-red (see below); hyphae interwoven, but 

becoming vertically orientated towards the hymenium, hyaline, c. 1-7-2 fxm wide; ascogenous 

hyphae short-celled, to 5 /xm wide, contents sometimes with dilute orange, K+ purple-red 

pigment. Excipulum reflexed to below the surface of surrounding thallus, thin, dilute brownish 

or ± hyaline in places, sometimes darkish brown at outer edge; hyphae radiating, branched and 

anastomosing, c. 1-5-2 /xm wide. 

Pycn/rf/fl not seen. 

Chemistry: Thallus surface and sections K— , C-, KC— , PD-, 

not analysed by t. I.e. 

I-; apothecia sections C— 

Observations: This species is placed in Micarea with some hesitation, owing to its unusual 

habitat (for a Micarea) and large-celled, thick-walled, non-micareoid phycobiont. However, 

much the same habitat is shared by M. subnigrata (a species with a typical micareoid phycobiont) 

and no other fungal characters appear to merit its exclusion. It is similar in appearance to M. 

subnigrata which can be distinguished by its brown epithecium and usual presence of pycnidia 

containing helicoid macroconidia. Superficially, M. intrusa is virtually indistinguishable from 

Table 5 Species associated with Micarea intrusa in Norway (see text). 

Aspicilia morioides 

Caloplaca of. lamprocheila 

Candelariella vitellina 

Cornicularia normoerica 

Fuscidea cyathoides 

F. intercincta 

F. tenebrica 

Haematomma ochroleucum 

Lecanora badia 

L. gangaleoides 

L. intricata 

L. polytropa 

L. cf . salina 

Lecidea fuliginosa 

L. fuscoatra 

L. lactea 

L. lapicida 

L. leucophaea 

L. vorticosa 

Micarea subnigrata 

Opegrapha gyrocarpa 

Pertusaria dealbescens 

P. lactea 

Placopsis gelida 

Porina chlorotica 

P. guentheri 

Ramalina subfarinacea 

Rhizocarpon geographicum 

R. lecanorinum 

R. riparium subsp. lindsayanum 

Rinodina gennarii 

Schaereria tenebrosa 

Stereocaulon vesuvianum 

Trapelia involuta 

Umbilicaria spodochroa 

± vertical 

rocks 

horizontal 

rocks 

;


forms of Scoliciosporum umbrinum with a well developed thallus, but that species has 

vermiform or sigmoid-curved spores. The phycobiont in M. intrusa and S. umbrinum appears to 

be the same; any future critical appraisal of the delimitation oi Scoliciosporum should seriously 

consider the generic disposition of M. intrusa. 

The pale orange, K+ purple-red pigment commonly found in the ascogenous hyphae, asci and 

spores has not been seen by me in any other Micarea, although it is conceivable that it is the 

same, or similar, to the pigment found in the goniocysts of M. hedlundii {q. v.). 

Habitat and distribution: M. intrusa occurs on hard siliceous (igneous) rocks in communities 

referable to the Umbilicarietum cylindricae in its broad sense (James etal., 1977). In Nylander's 

protologue of Lecidea aphanoides it is said to occur on calcareous rock, but this is erroneous; 

application of 50% HNO3 to fragments of substratum from the holotype produced no efferves- 

cence. The formation of small patches amongst other crustose lichens indicates that M. intrusa 

may be, at least facultatively, parasitic. Such suggestions have been previously propounded by 

Magnusson (1942), Poelt (1958), and Wirth (1973), all of whom related its behaviour to that of 

Lecanora intrudens and Lecidea furvella. In several collections, including the type of Lecidea 

intrusa, it occurs amongst the areolae of Huilia panaeola, but it is by no means confined to that 

'host'. 

From her studies on the island of Sotra in Hordaland, Norway, Miss L. Skjolddal informs me 

that M. intrusa occurs on sheltered, sunny, west or south-west facing exposures of gneiss and 

amphibolite, on steep surfaces or, in one case, on a ± horizontal surface. Miss Skjolddal kindly 

provided me with a list of associated species (Table 5). 

M. intrusa is probably widespread in areas of western Europe with exposed, hard, igneous 

rocks; however, it is a very inconspicuous species and records are wanting from many potential 

localities. I have seen material from southern Scandinavia (several localities). North Norway 

(Finnmark), and the Grampian Mountains of Scotland. In addition, Wirth (1973) cites collections 

from France (Vosges) and West Germany (Schwarzwald). 

Exsiccata: Norrlin & Nyl. Herb. Lich. Fenn. 182 (BM). 

18. Micarea leprosula (Th. Fr.) Coppins & A. Fletcher 

(Figs 18, 53-54; Map 8) 

in Fletcher in Lichenologist 7: 111 (1975). - Bilimbia milliaria y leprosula Th. Fr. , Lich. Scand. 2: 382 

(1874). - Micarea violacea var. leprosula (Th. Fr.) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 

(3): 81, 92 (1892). - Bilimbia leprosula (Th. Fr.) H. Olivier in Bull. Geogr. bot. 21: 179 (1911). - Bacidia 

leprosula (Th. Fr.) Lettau in Hedwigia 52: 133 (1912). Type: Sweden, Uppsala, Witulfsberg [Wittuls- 

berg], 26 vi 1852, Th. M. Fries (UPS - lectotype! [t.l.c. : argopsin and gyrophoric acid]). 

Thallus effuse, superficial, of convex to subglobose areolae c. 100-350(-400) fxm diam; these 

usually coalesce toward the centre of the thallus, whence they may proliferate, producing 

secondary granular-areolae, such that the thallus becomes thicker (to c. 700 /xm thick). Areolae 

blue-grey or, more rarely, grey-brown, matt with a minutely roughened surface with white flecks 

(x50 lens); lower sides of ± globose areolae, and areolae in shaded situations, often unpigmented 

and greenish white. Areolae fragile and often breaking down or eroding to form 

sorediate patches with irregularly shaped soredial granules c. 20-50 /xm diam. Sections of intact 

areolae ecorticate and without an amorphous hyaline covering layer; outermost hyphae often 

coloured with greenish or brownish pigment, K— , HNO3-I- reddish. Phycobiont micareoid, cells 

4-7 /xm diam. 

Apothecia often absent, at first adnate but often becoming constricted below, convex to 

subglobose, commonly tuberculate, immarginate or rarely very faintly marginate when young, 

matt or slightly glossy, dark blue-grey or black, 0- 15-0-4 mm diam, or to 0-7 mm diam when 

HNO3-I- 

tuberculate. Hymenium 45-60 /xm tall, hyaline or dilute green, but dark green (K— , 

red) in upper part (epithecium). Asci clavate, 40-55x14-16 /xm. Spores oblong-ellipsoid, 

oblong-fusiform or clavate-fusiform, often curved, (l-)3-septate, 14-26(-29)x4-5(-5-5) /xm. 

Paraphyses numerous, branched and anastomosing, (0-7-)l-l-5 /xm wide; apices often more


richly branched, not or only slightly incrassate (to 1-8 /xm wide) and without closely adhering 

pigment (epithecial pigment confined to surrounding gel-matrix). Hypothecium c. 35-45 txm 

tall, but becoming much taller in markedly convex or tuberculate apothecia, dilute straw- or 

fuscous-brown, pigment confined to gel matrix; hyphae interwoven, but becoming outwardly 

orientated towards the hymenium and excipulum, c. 0-8-1-5 /u,m wide; ascogenous hyphae with 

swollen cells, c. 2-6 /am wide. Excipulum distinct in young, moderately convex apothecia, but 

reflexed and obscured in subglobose or tuberculate apothecia, dilute fuscous brown (K— ); 

hyphae radiating, branched and anastomosing, c. 0-5-1 -5 /u,m wide. 


Chemistry: Thallus K-, C+ red, PD-I- red; apothecia in section C-l- red; t.l.c: argopsin and 

gyrophoric acid. 

Observations: M. leprosula is characterized by its sterile or sparingly fertile, granular thallus 

composed of blue-grey (or brown-grey), fragile areolae which often dissolve (or become 

abraded) to form contrasting, yellowish green, sorediose patches. These characters are shared 

by the much rarer M. subleprosula, and the thalli of both react C+ red. However, M. 

subleprosula can be separated by the PD-I- yellow reaction of its thallus due to the presence of 

alectorialic acid. When fertile M. subleprosula is found to have much larger, mostly 7-septate 

spores. M. leprosula is usually distinctive in the field (although material should always be 

checked with M. subleprosula in mind) but can easily be overlooked when it occurs on 

bryophytes on shaded rocks or trees, whence the thallus often lacks its characteristic bluish tinge 

and the areolae are thinly scattered. The species is usually sterile but fertile specimens are 

occasionally encountered, especially in sheltered (but not deeply shaded) situations, such as on 

boulders and old walls in woodland. 

Hedlund (1892) regarded M. leprosula as a variety of M. peliocarpa (q.v.) and the apothecia 

of the two species are somewhat similar. However, those of M. leprosula exhibit several minor, 

yet significant, differences; spores tend to be slightly longer; apothecia tend to be more 

markedly convex and more frequently tuberculate; hypothecium and excipulum are dilutely 

pigmented with a dull brown pigment; and paraphyses never become thickened and pigmented 

at their apices. The two species also differ in several fundamental aspects of thallus morphology 

and chemistry. With regard to the production of argopsin and to the presence of a dull brown 

pigment in the excipulum and hypothecium M. leprosula shows some affinity to M. lignaria var. 

lignaria, with which it often occurs. However, M. lignaria lacks gyrophoric acid, has usually 

longer and more septate spores, and somewhat stouter and only sparingly branched paraphyses. 

The areolae of M. leprosula and M. subleprosula differ from those of such species as M. lignaria, 

M. peliocarpa, and M. cinerea, in lacking an amorphous hyaline covering layer (see 'morpho- 

logy')- 

Habitat and distribution: M. leprosula is widely distributed in upland areas in the north and 

west of Britain, occurring at altitudes from about sea-level to at least 1100 m. It is found, usually 

on moribund bryophytes (e.g. Andraea spp. and Rhacomitrium spp.) on rocky ledges, boulders, 

and old walls, in exposed hilly areas, mountain sides or woodlands. Associated lichens noted 

amongst British collections include Arthrorhaphis citrinella, Baeomyces rufus, Belonia incar- 

nata, Cladonia cervicornis, C. coccifera, C. crispata, C. portentosa, C. squamosa, C. subcervicornis, 

Coelocaulon aculeatum, Lecidea granulosa, L. icmalea, Lepraria neglecta, Micarea 

lignaria, M. peliocarpa, and Vorarlbergia renitens. From Scotland I have seen two corticolous 

specimens, on trunks of Alnus and Betula; associated species present included Cladonia 

coniocraea, C. squamosa, Lecidea icmalea, Micarea melaena, and Platismatia glauca. 

There are two outlying localities in the lowlands of southern England. In Dorset it occurs on 

waste heaps of slag clay with M. lignaria (q. v.); and in Kent it was found with M. peliocarpa on 

wood chips lying at the side of a woodland track in a chestnut (Castanea) coppice. 

M. leprosula is poorly recorded outside Britain, probably because it is so often sterile. To 

date, I have seen one specimen from Norway (Hordaland) and several collections from mid- and 

southern Sweden. Further south it seems to be well represented in the Alps, and I have seen one


Map 8 Micarea leprosula # 1950 onwards O Before 1950 

collection from Bohemia (Czechoslovakia). This Bohemian record, and another from the Black 

Forest (Schwarzwald) were made from exsiccate specimens {Kutdk 417 and Migula 1) of M. 

lignaria growing on wooden roof shingles. 

Exsiccata: Kutak Lich. Bohem. 417 p. min. p. (0). Magnusson Lich. Sel. Scand. 208 (BM, C, GZU). 

Migula Crypt. Germ. \p. min. p. (C, E, M). 

19a. Micarea lignaria (Ach.) Hedl. var. lignaria 

(Figs lA, 3D-E, 19, 45, 53-54; Map 9) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 82, 93 (1892). - Lecidea lignaria Ach. in K. svenska 

VetenskAkad. Handl. 1808: 236 (1808). - Bilimbia lignaria (Ach.) Massal. , Ric. Lich. Crost. : 121 (1852). 

- Bacidia lignaria (Ach.) Lettau in Hedwigia 52: 132 (1912). Type: Sweden: 'Svecia', on lignum, 

(H-ACH 265 - lectotype; BM - ? isotype!). 

Lecidea milliaria Fr. in K. svenska VetenskAkad. Handl. 1822: 255 (1822). Type: Sweden, Smaland, 

Femsjo, on lignum, E. M. Fries (UPS - lectotype! [t.l.c: argopsin]). 

Lecidea geomaea Taylor in Mackay, Fl. Hib. 2: VIA (1836). Type: Ireland, South Kerry, Dunkerron, T. 

Taylor (BM - lectotype!; BM ex K - isolectotype!). 

Bilimbia milliaria a. [var.] lignaria** [f.] calamophila Korber, Parerga lich. : 171 (1860). Type: Germany: 

'an Schilfrohrdachern um Miinster', J. G. F. X. Lahm (L 910, 144-1685 - lectotype!; L 910, 144-1689 - 

isolectotype! [t.l.c: argopsin]. 

Bilimbia milliaria (3. [var.] saxicola Korber, Parerga lich. : 171 (1860). Type: Germany: 'Bilimbia trochiscus 

Korb. prim, B. miliaria p. saxicola Kb.! Extersteine', on sandstone, ? Beckhaus (WRSL- lectotype!).


Paratype: Germany: 'Ibbenbiiren in Westfalen, Lahm, Hb. Hepp.', on sandstone (BM!). 

Stereocauliscum gomphillaceum Nyl. in Flora, Jena, 48: 211 (1865). - Micarea gomphillacea (Nyl.) Vezda 

in Folia geobot. phytotax., Praha 5: 321 (1970). - Bilimbia milliaria f. [as 'f?'] gomphillacea (Nyl.) 

Blomb. & Forss., Enum. PI. Scand.: 82 (1880). - Micarea lignaria [as 'ligniaria'] f. gomphillacea (Nyl.) 

Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 94 (1892). Type: Finland, Tavastia australis, 

HoUola, Tiirismaa, Pirunpesa, [on granite rock], 1863,/. P. Norrlin (H-NYL 40217 -lectotype! [t.l.c: 

argopsin];isolectotypes: H-NYL 40214!, 402151,40216!, 40218! and p.m. 631!). 

Lecidea sabuletorum var. milliaria f. nigrata Nyl. in Not. Sdllsk. Fauna Fl. fenn. Forh. 8: 151 (1866). Type: 

USSR, Kola Peninsular, Lapponiaponojensis,Ponoi, 1863, M /. Fe/Zman (H-holotype!;H-isotype!). 

See note (i) below. 

Lecidea milliaria var. triseptata Nyl. in Lamy in Bull. Soc. bot. Fr. 25: 441 (1878). - Bilimbia triseptata 

(Nyl.) Arnold in Flora, Jena 67: 572 (1884), non Bacidia triseptata (Hepp in ZoU.) Zahlbr. Type: France: 

Haute Vienne, 'Sur des roches entre Chalard et St Junien, Lamy (H-NYL 18367! - lectotype, as 'L. 

milliaria var. 3-septata Nyl.'). See note (ii) below. 

Lecidea meizospora Harm, in Bull. Seanc. Soc. Sci. Nancy II, 33 (1898): 63 (1899). - Bacidia meizospora 

(Harm.) Zahlbr., Cat. lich. univ. 4: 122 (1926). Type: France, Vosges, Docelles, on Calluna roots, V. 

and H. Claudel & F. J. Harmand (ANGUC - lectotype!). 

Lecidea trisepta var. polytropoides Vainio in Ark. Bot. 8 (4): 106 (1909). Type: USSR, Magadanskya 

Oblast (N.E. Siberia), 'Pitlekai, peninsula Jinretlen' [c. 67°7'N 173°24'W], on plant debris, 1878-9, E. 

Almquist (TUR-VAINIO 21274, fertile part- lectotype!). See note (iii) below. 

Patellaria milliaria var. xylophila Wallr., Fl. crypt. Germ. 1: 349 (1831); nom. inval. (Art. 26). 

Lecidea milliaria var. saxigena Leighton, Lich. Brit. Exs. 210 (1856); nom. nudum (Art. 32); see note 

under M. peliocarpa. Type: England, Shropshire, Neescliff [Nesscliff] Hill, Leighton, Lich. Brit. Exs. 

210 (E- lectotype!; isolectotypes: BM! DBN! MANCH! NMW!). 

Notes: (i) The type material mainly consists of the black stroma of Bryomyces gymnomitrii Dobb. on 

Gymnomitrium sp. , with only a few apothecia of M. lignaria present. 

(ii) Lamy (loc. cit.) gives the locality as 'entre le Chalard et Saint-Yrieix'. The label on the lectotype 

specimen is entirely in Nylander's handwriting and there may have been an error in transcription. 

(iii) Vainio's diagnosis included the description of soralia. A small portion of the type specimen is a 

sterile sorediate thallus (C- , PD- 

phycobiont cells 7-14 /am diam) but it is not a Micarea and is not part of 

; 

the larger, fertile portion. The latter belongs to Micarea lignaria and is chosen as lectotype. 

Thallus effuse, partially endoxylic or endocuticular, but more usually developed superficially 

as convex to ± globose areolae. Areolae c. 80-250(-300) /xm diam, discrete, scattered to 

contiguous, sometimes coalescing to form an uneven crust, grey-white and ± glossy to 

grey-green or bluish grey and matt, sometimes brown-grey due to the ramification through their 

surface of brown, toruloid hyphae (cells c. 5-7x4-5-7 /xm) belonging to an unknown fungus. 

Areolae in section, with a hyaline amorphous covering layer (c. 4—12 /xm tall); walls of hyphae in 

outermost (exposed) parts often blue-green (K-, HNO3+ red); algal layer usually in direct 

contact with substratum, but in larger areolae the algal layer may be c. 90-135 ixva tall lying 

above a medulla. Medulla, when present, up to c. 100 îm tall, consisting of loosely interwoven 

hyphae c. 1-1-5 /xm wide, intermixed with dead algal cells and debris derived from the 

substratum. Phycobiont mic2iVQO\d, cells c. A-1 /xm diam. 

Apothecia scattered, more usually numerous and crowded, often confluent, immarginate, 

convex-hemispherical, soon becoming constricted at the base and hence ± globose, occasionally 

becoming stipitate with a grey-black 'stipe' up to 1 mm tall ('f. gomphillacea''), black, matt or 

slightly glossy, rarely blue-grey (shade forms), 0-15-0-6(-0-9) mm diam. Hymenium 50-75 /xm, 

often not sharply delimited from the hypothecium, dilute olivaceous or aeruginose-green (K- 

HNO3-I- red) in upper, and sometimes lower, part(s) (the middle part often being ± hyaline); in 

the uppermost part (epithecium) the pigment is more dense and is present not only in the gel 

matrix but also in the walls of the paraphyses; in reflexed parts of the hymenium towards the base 

of the apothecium the green pigment is often replaced by a dilute brown pigment; minute (less 

than 2x1 yu,m) granules of violet (K+ intense aeruginose) pigment are sometimes present and 

lightly scattered through the hymenium. Asci clavate, 45-50x11-19 /xm. Spores fusiform, 

straight or slightly curved, (0-)3-7 septate, 16-36(-38)x4-6(-7) /xm. Paraphyses numerous, 

simple or slightly branched above, but more richly branched in the transition zone between the 

hymenium and excipulum, sometimes anastomosing, 1-3-1-8 /xm wide; apices often slightly 

,


incrassate and coloured with green or greenish brown pigment and then up to 3 /xm wide, usually 

coherent (even in K) and, together with the surrounding pigmented matrix, form an epithecium. 

Hypothecium c. 100-230(-350) /xm tall, dilutely pigmented with pigment confined to gel matrix; 

upper part dull aeruginose or olive-brownish K- or + dulling, HNO3+ reddish); lower part 

often without greenish tinge, and then being dilute brownish or ± hyaline; hyphae interwoven or 

some ± vertically orientated near the hymenium, c. 1-1-7 /am wide; ascogenous hyphae with 

short, swollen cells c. 2-5-4 /xm wide. Excipulum ± distinct in sections of young, hemispherical 

apothecia, but soon becoming strongly reflexed and not sharply delimited from the hymenium, 

dilute brown or sometimes darkish brown along the outer edge. Hypothecial and excipular 

tissues sometimes elongated vertically to form a stipe ('f. gomphillaced') 

Pycnidia inconspicuous, ± immersed, with walls green (K-, HNO3-I- red) in exposed upper 

parts and ± hyaline in immersed lower parts; of three types [only type (c) is common]: (a) c. 100 

ixm diam; conidia {macroconidia) curved or hamate, 0-3-septate, 16-22 Xc. 1 /xm; (b) c. 100-140 

/xm diam; conidia (mesoconidia) ± cylindrical, oblong-ellipsoid, obovoid or oblong-obovoid, 

usually distinctly truncate at proximal end, often 2-3-guttulate, 4-7(-7-6)xl-2-l-8 fxm; (c) c. 

40-50 îm diam; conidia (microconidia) narrowly cylindrical, only faintly truncated at proximal 

end, eguttulate, (4-5-)5-7(-8)x 0-8-1 /xm. 

Chemistry: Thallus K-, C-, KC-, PD-I- red; sections of apothecia C-, PD-; t.l.c: 

argopsin. 

Observations: Micarea lignaria is characterized by its whitish to grey, convex to ± globose 

areolae which are PD+ red (argopsin; but see var. endoleuca), black, markedly convex to ± 

globose apothecia, green upper hymenium, dilute greenish or dilute olive-brownish hypothecium, 

'^nd 3-7-septate, fusiform spores. When on lignum the thallus is often reduced to small 

scattered areolae and sometimes it is ± entirely endoxylic. The apothecia are less variable in 

appearance, and pale (shade) forms are very rare. However, the apothecia are occasionally 

found (especially on rock in dry underhangs) to be stipitate with a 'stipe' (composed of vertically 

extended hypothecial and excipular tissues) up to 0-4 mm, or even 1 mm, tall (Fig. 3E). These 

forms have been ascribed the status of form, variety, species and even genus, viz.: Nylander's 

new species and genus Stereocauliscum gomphillaceum. In the type material of this, some 

'stipes' are extremely tall (up to 1 mm) and composed of vertically proliferating apothecia, with 

the apical apothecia being the youngest; the apothecia are immature (or arrested in their 

development) with few asci and spores. In my opinion these stalked forms of A/, lignaria result 

from abnormal development in response to adverse environmental conditions and are, there- 

fore, not worthy of taxonomic recognition at any rank. 

M. lignaria is often confused with M. cinerea and M. peliocarpa, but these two species have 

usually more flattened and often paler apothecia, more richly branched paraphyses, and a 

hyaline hypothecium. In addition, they contain gyrophoric acid, resulting in the C-l- orange-red 

(quickly fading) , PD 

— reactions of their thalli and apothecia. M. ternaria {q. v. ) is very similar to 

M. lignaria, but has more flattened apothecia with a more discernible excipulum and spores 

which are never more than 3-septate; furthermore, it lacks any lichen substances. 

Habitat and distribution: M. lignaria occurs on a wide range of substrata, but is most common 

in upland districts, growing over bryophytes or plant debris on siliceous rocks and walls, or on 

exposed peaty ground. Associated lichens found in such habitats in Britain include 

Arthrorhaphis citrinella, Baeomyces rufus, Cladonia coccifera, C. floerkeana, C. squamosa, C. 

subcervicornis, Coelocaulon aculeatum s. lat., Lecidea granulosa, L. icmalea, Lecidoma demissum, 

Lepraria neglecta, Micarea leprosula, M. peliocarpa, Ochrolechia frigida, Parmelia 

saxatilis, and Pseudephebe pubescens. It often grows over bryophyte mats that are heavily 

invaded by gelatinous algae, and in such situations in Scotland it has been found with the rare, or 

overlooked, Arctomia delicatula and Belonia incarnata. When growing directly on rock M. 

lignaria is mostly confined to sheltered, shaded situations and is sometimes present in the 

Micareetum sylvicolae in rock underhangs. It is usually present in abundance in the old lead and 

zinc mine workings of the Pennines and Scotland, where it grows over decaying bryophytes and 

.


Map 9 Micarea lignaria var. lignaria # 1950 onwards O Before 1950 

plant debris, loose stones, pieces of timber and sack-cloth. Mr V. Giavarini has recently found it 

in Dorset, growing on waste heaps of slag clay, in the company of M. leprosula, Baeomyces 

roseus, Cladonia arbuscula, C. ciliata, C. furcata, C. portentosa, and Coeldcaulon aculeatum s. 

lat. M. lignaria is very rare in the lowlands of south-east England, but has been found there 

growing directly on the rock of east or north facing sandstone walls and churchyard memorials, 

and also on natural sandstone outcrops of the Sussex Weald. 

In upland areas M. lignaria is frequently found on the exposed lignum of fallen trunks 

(especially of conifers) and old timberwork, with, for example, Cladonia spp., Hypogymnia 

physodes, Micarea peliocarpa, Mycoblastus sterilis, Ochrolechia turneri, Parmelia saxatilis, and 

Lecanora polytropa (worked timber) . Occurrences of M. lignaria on the bark of healthy trees 

are rather rare, and in Britain are confined to the high rainfall areas of the north and west, where 

it has been collected on Alnus, Betula, Fraxinus, Ilex, Quercus, and old Sambucus. 

Reports of M. lignaria on mosses on limestone rocks usually result from the misidentification 

of Bacidia sabuletorum or Toninia lobulata. However, on a few occasions I have seen M. lignaria 

growing on thick bryophyte cushions or mats at high altitudes over limestone in the north 

Pennines, and over calcareous mica-schist in the Breadalbane Mountains; in such situations the 

pH and calcium content of the substratum is presumably kept low by the leaching effect of the' 

high rainfall in those areas. 

The altitude range of M. lignaria in Britain is from sea-level to about 1200 m (Ben Lawers), 

although it may well occur at higher altitudes in the Ben Nevis group and the Cairngorni 

Mountains. Higher altitudes are attained in the mountains of central Europe, from where it has

146 


been collected at 2800 m in the Austrian Tirol, c. 2000 m in the Tatry mountains of Poland and 

Czechoslovakia, and at nearly 2500 m in the Transylvanian Alps of Romania. 

M. Ugnaria is common in northern and western Britain, but is very rare, with only a few 

scattered records, in lowland, southern England east of Devon. It is widely distributed 

throughout most of Europe, although in southern Europe it appears to be confined to the 

mountains: like most species of Micarea it seems to avoid the lowland Mediterranean regions. 

Records suggest that it is rare in arctic Fennoscandia, although I have seen material from north 

Finland and the Kola Peninsula. It has been found in the Azores at altitudes of about 900 m but I 

do not know of it from elsewhere in Macaronesia. From outside Europe I have seen material 

from the north coast of western Siberia, the Franconia Mountains (at c. 1450 m) of New 

Hampshire in the USA, and from Brazil (Serra Montiqueira, at c. 1900 m). 

Exsiccata: Anzi Lich. Lang. 148 (BM). Arnold Lich. Exs. 348A, B (BM ex K, L, M). Bohler Lich. Brit. 

85 (E). Claudel & Harm. Lich. Gall. 43 (BM). Cumm. Dec. N. Am. Lich. ed. I: 302 (BM, C), ed. 2: 232 

(M). Flotow Lich. Exs. 129A, E and 131 (UPS). Fries Lich. Suec. 29 (E, M, UPS); 98 (E). Harm. Guide 91 

(UPS). Harm. Lich. Loth. 852 (M). Hav. Lich. Norv. 555 (C). Hertel Lecid. 34 (E, GZU). Johnson Lich. 

Herb. 453 (E). Kalb Lich. neotropici 22, 186 (hb Kalb). Krypt. Exs. Vindob. 658 (BM, BM ex K, C, M). 

Kutak Lich. Bohem. 417 (O). Larb. Lich. Caes. Sarg. 83 (BM). Larb. Lich. Herb. 272 (BM). Leighton 

Lich. Brit. 210 (BM, DBN, E, MANCH, NMW), 238 p.p. (BON, DBN, E, M, MANCH), 386 (BM, 

BON, DBN, E, FRS, M). Lojka Lich. Hung. 61 (BM, BM ex K, M). Malme Lich. Suec. 288 (C, M, S). 

Migula Crypt. Germ. Ip.p. (BM, C, M, MANCH); 226 (BM, C, MANCH). Moug. & Nestl. Stirp. Crypt. 

1430 (E). Mudd Lich. Brit. 156 (BM, E, M, MANCH), 157 (E, MANCH), 158 (BM, E, M, MANCH). 

Norrlin & Nyl. Herb. Lich. Fenn. 319A, B (BM, C, H). Oliv. Lich. Orne 344 (M, S). Pisut Lich. Slov. 156 

(BM, M). Rabenh. Lich. Eur. 322, 603 (BM, M). Roum. Lich. Gall. 193 (BM), 232 (BM, M). Samp. Lich. 

Port. 147 (LD). Schaerer Lich. Helv. 196p.p. (BM, BM ex K, E). Vezda Lich. Bohem. 133, 258 (LD, M). 

Vezda Lich. Sel. 516, 858, 1036, 1088 (BM). Zwackh. Lich. Exs. Ill (BM). 

19b. Micarea lignaria var. endoleuca (Leighton) Coppins, comb. nov. 

(Figs 53-54; Map 10) 

Lecidea milliaria var. endoleuca Leighton, Lich. Fl. Brit., edn 3: 363 (1879). Type: Ireland, West Galway, 

ontheDoughraugh, 1875, C. Lar/?fl/e5îeA-(BMexK-lectotype!; BM-? isolectotype!). 

Thallus identical to var. lignaria except that the areolae usually have dull yellowish 

(isabelline) tinge. 

Apothecia and pycnidia (containing microconidia) identical to var. lignaria; pycnidia containing 

macroconidia or mesoconidia not known. 

Chemistry: Thallus K- or Kf+ yellow, C+ yellow-orange (persistent), KC-I- reddish orange 

(persistent), PD-; sections of apothecia C-; t.l.c: xanthone(s), possibly arthothelin and 

thiophaninic acid. 

Observations (including habitat and distribution) : My 

preliminary studies on M. lignaria found 

most specimens to have a C-, PD+ red thallus reaction, but a few otherwise ± identical 

specimens from western Ireland, north Wales, and western Scotland gave C+ persistent orange, 

PD- reactions. Subsequent t.l.c. analysis proved the normal form to have the then unidentified, 

P+ red compound found also in Phyllopsora rosei (Coppins & James, 1979), and the anomalous 

form to contain two xanthones (possibly arthothelin and thiophaninic acid). Xanthones often 

impart a yellowish tinge to lichen thalli (e.g. Lecidella elaeochroma, L. scabra, Pertusaria 

flavida, and P. hymenea) and this is the case for the xanthone containing race of M. lignaria; the 

colour difference is most evident when the two races occur together. Of all the types of names 

attributable to M. lignaria s. ampL, only that applicable to the 'var. endoleuca Leighton' was 

found to contain xanthones, and that name is here adopted at the same rank. Although the var. 

endoleuca is morphologically identical to the var. lignaria it has a sympatric but much more 

restricted geographical range. In the British Isles it is confined to the more oceanic districts with 

one rather anomalous occurrence in Surrey (Leith Hill). Furthermore, it appears to have a lower 

altitudinal range, with a collection at 600 m in Snowdonia representing its known upper limit in


Map 10 Micarea lignaria var. endoleuca # 1950 onwards O Before 1950 

the British Isles. Only four examples of var. endoleuca have been discovered from outside 

Britain: from Baden (Heidelberg) and Bavaria (Bayreuth) in Germany, Bern (Langnau) in 

Switzerland, and Trentino (Appiano) in northern Italy. It should be checked for amongst 

material from other regions, especially south-west Norway, Bretagne (France) and the Pyre- 

nees. 

I believe that the var. endoleuca should be retained at varietal rank. However, as the 

depsidone argopsin (I'-chloropannarin) of var. lignaria is probably biogenetically distinct from 

the xanthones of var. endoleuca, a case could be made for its recognition at species rank. A 

comparative example is Parmeliopsis ambigua versus P. hyperopia (see Hawksworth, 1976). 

20. Micarea lithinella (Ny 1 ) Hedl 

. 

(Figs4A,20A-B;Mapll) 

. 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 97 (1892). - Lecidea lithinella Nyl. in Flora, Jena 

63: 390 (1880). - Lecidea lithinella Nyl. in Flora, Jena 45: 464 (1862); nom. nudum (Art. 32). Type: 

Germany, Bayern, 'Sandsteineblocke am Waldwege von Banz nach Altenbanz in Oberfranken', ix 

1860, F. Arnold 957 (M - lectotype!; H-NYL 19192 - isolectotype (fragment)!). 

Thallus effuse, saxicolous, often a thin indistinct crust between protruding grains of rock, but 

somes developing irregularly rounded, convex, more rarely subglobose, whitish areolae, c. 

40-100 /xm diam. Phycobiont micareoid, cells 4—6-5 /xm diam. 

Apothecia numerous, immarginate, adnate, convex, apparently never tuberculate, pallid or


dull yellow-orange to reddish brown (never black), (O-l-)O- 15-0-4 mm diam. Hymenium 35-50 

îm tall, hyaline but often tinged dilute straw in upper part. Asci cylindrical-elavate, 35-50x8-13 

/xm. Spores ovoid or fusiform-ellipsoid, simple, 6-5-9-5X 2-8-4 ^tm. Paraphyses scanty, simple 

or irregularly forked above, 0-7-1 ^tm wide but sometimes gradually widening towards the 

apices and up to 1-5 //-m wide, hyaline throughout. Hypothecium c. 50-70 fxm tall, yellow-straw 

to dilute orange-brown, K- , HNO3- 

pigment confined to gel-matrix , and hyphal walls hyahne. 

Excipulum absent (not detected even in sections of young apothecia). 

Pycnidia apparently rare, very inconspicuous, immersed, c. 40 /am diam, with hyaline walls. 

Conidia (microconidia) narrowly cylindrical, 4-5-7x0-8-1 fim. 

Chemistry: Thallus C— , K— 

, PD-; section of apothecia C-; t.l.c: not tested. 

Observations: M. lithinella exhibits little variation except that between, and sometimes 

within, collections there is an intergradation from pallid apothecia with a dull yellowish 

hypothecium to darker apothecia with an orange-brown hypothecium. This variation is correlated 

with differences in exposure to light. M. lithinella is easily confused with forms of M. 

bauschiana with pallid apothecia. Difficult specimens can be determined by a careful examination 

of the phycobiont, which is micareoid in M. lithinella but non-micareoid (cells c. 7-13 jxm 

diam) in M. bauschiana. M. lutulata has darker apothecia which are commonly tuberculate, a 

darker (opaque) hypothecium, an often greenish hymenium, and non-micareoid phycobiont. 

M. muhrii is normally lignicolous but has once been found on rocks; it has adnate apothecia like 

M. lithinella but they are usually darker and have a taller, dark reddish brown hypothecium, and 

usually a greenish upper hymenium. M. myriocarpa shares a similarly coloured hypothecium 

with M. lithinella, but has smaller, globose to tuberculate apothecia and smaller, 1-septate 

spores. 

In their keys to Micarea Vezda & Wirth (1976) and Poelt & Vezda (1977) state that M. 

lithinella has a K+ violet hymenium and often 1-septate spores; this is clearly an error, probably 

due to confusion with saxicolous forms of M. denigrata or M. prasina. 

Habitat and distribution: M. lithinella occurs on acidic rocks, especially hard sandstone. 

Associated species encountered on the specimens examined include Baeomyces rufus (parasi- 

tized by Thelocarpon lichenicola on the lectotype of Lecidea lithinella), Huilia crustulata 

Rhizocarpon obscuratum, Scoliciosporum umbrinum, Trapelia coarctata, and T. aff . obtegens. 

This list of associates, plus information on some of the packets, suggest that M. lithinella has 

mostly been found in humid situations on outcrops and boulders by woodland roads. It was 

collected by Arnold, Lahm and von Zwackh from several areas of Germany: Bayern, Nordrhein-Westfalen 

and Baden-Wiirttemberg, respectively. In Sweden it was collected in Uppland 

by Hedlund who (1892: 97) also cited a collection by Blomberg from Sodermanland. One British 

specimen has recently been found: South-east York (V.C. 61), Wharram Quarry Nature 

Reserve, 44/85. 65, 1969, Coppins (E). At this locality M. lithinella was growing with Scoliciosporum 

umbrinum on the underside of a large flint; the upper, more exposed, part of the flint 

was colonized by Lecidea erratica and Rhizocarpon obscuratum. 

Exsiccata: Arnold Lick. Exs. 836 (BM ex K, H-NYL 19190, M). Lojka Lick. Univ. 233 (M). Malme 

Lick. Suec. 125 (M, S). Zwackh Lich. Exs. 590 (H-NYL p.m. 5404, M). 

21. Micarea lutulata (Nyl.) Coppins 

(Figs 20c, 44B; Map 11) 

in D. Hawksw., P. James, & Coppins in Lichenologist 12: 107 (1980). - Lecidea lutulata Nyl. in Flora, 

Jena56: 297 (1873). Type: Jersey, Rozel meadow, bases of rocks, 1873, C. Larbalestier (H-NYL 10696lectotype!; 

BM-isolectotype!; M- probable isolectotype!). See note below. 

Lecidea laxulaNyl. in Flora, Jena 5S: 11 (1875). Type: Finland, Tavastiaaustraiis,Luhanka, Hietala, 1874, 

E. A. Lang [Vainio] 303 (H-NYL 20689 - lectotype!; H - isolectotype!). 

Lecidea poliodes Nyl. in Flora, Jena 58: 10 (1875). - Micarea poliodes (Nyl.) Vezda in Vezda & V. Wirth in 

Folia geobot. phytotax., Praha 11: 99 (1976). Type: Finland, Tavastia australis, Evo, [on schistose rock 

with M. sylvicola],J. P. Norrlin (H-NYL 20683 - holotype!).


Lecidea antrophila Larb. ex Leighton in Trans. Linn. Soc. London (Bot.) II, 1: 242 (1876). Type: Ireland, 

West Galway, Mwellan near Kylemore, in the interior of a cave, 1877, C. Larbalestier (BM ex K - 

lectotype!; isolectotypes: BM!, BM ex K!). 

Lecidea paucula Nyl. in Flora, Jena 59: 573 (1876). Type: Ireland, West Galway, 'Montagnes de Maam' 

[Maumturk Mountains], iii 1876, C. Lflr/?fl/e5ner(H-NYL 20090 -holotype!). 

Micarea umbrosa Vezda & V. Wirth in Folia geobot. phytotax., Praha 11: 93 (1976). Type: Germany, 

Baden, Siidschwarzwald, between Neuhausle and Altglashiitte near St. Margen, 830 m, 2 v 1969, V. 

Wirth (hb. Wirth 1609 - holotype!). 

Lecidea granvina Vainio in Havaas in Bergens Mus. Arb. 1909 (1): 31 (1909); nom. nudum (Art. 32). Spec. 

orig.: Norway, Hordaland, Hardanger, Granvin, 10 xi 1900,7. J. Havaas (BG!). 

Lecidea demarginata auct. p.p., non Nyl.; see 'Excluded Taxa'. 

Note: typification of Lecidea lutulata. Larbalestier's gatherings in BM and H, that were collected from 

Rozel meadow in 1873 and subsequently labelled 'Lecidea lutulata', consist of M. bauschiana, M. lutulata 

and M. peliocarpa. The specimen H-NYL 10696 contains M. lutulata only and agrees with the original 

diagnosis, e.g. 'hypothecium fusconigricans crassum', and is therefore selected as lectotype. 

Thallus effuse, thin (up to 40 /xm thick), ± smooth or finely rimose, usually becoming thicker 

(up to 600 jxm thick) and scurfy-granular (but never forming discrete areolae or goniocysts) pale 

, 

greenish grey or grey-green, sometimes straw-coloured in part, sometimes oxydated (on 

ferruginous rocks); in section ecorticate and without an amorphous hyaline covering layer. 

Phycobiont not micareoid; cells thin-walled, ± globose, c. 5-12 /xm diam, or ellipsoid and up to 

15x8/xm. 

Apothecia numerous, convex-hemispherical and immarginate from the start, later becoming 

± globose or tuberculate, grey-brown, dark brown or blackish, always turning blackish when 

moistened, 0-2-0-4 mm diam, or up to 0-8 mm diam when tuberculate. Hymenium 30-40 îm 

tall; upper part varying from hyaline or pale fuscous-brown through olivaceous to dark 

aeruginose, pigment often in patches corresponding to clustered apices of stout paraphyses, 

K- , HNO3+ red; lower part hyaline or dilutely coloured. Asci cylindrical-clavate, c. 30-40x7- 

10 /am. Spores ellipsoid, ovoid, or sometimes dacryoid, 6-8 (-9) x 2-3 (-3 -4) yam. Paraphyses 

rather scanty, of two types: p.p. evenly distributed, irregularly flexuose, simple, or forked or 

with short branches (especially in the upper part), sometimes anastomosing, 0-8-l(-l-5) ^tm 

wide, apices sometimes irregularly swollen to c. 2 ^tm wide; p.p. fasciculate, mostly simple, 

stout, c. 1-5-2 /xm wide, with swollen (to 3 /xm), coherent, pigmented apices. Hypothecium 

120-360 /x,m tall, dark and opaque, fuscous- or reddish brown, K- or -I- reddish intensifying (but 

never with purple or greenish tinges), HNO3-; hyphae interwoven, but becoming vertically 

orientated towards the hymenium, c. 1-5-2 /xm wide but coated with dense brown pigment and 

appearing 3-4 /xm wide; ascogenous hyphae with swollen cells c. 2-4 /xm wide. Excipulum not 

seen even in sections of young apothecia; the hymenium soon becomes reflexed so as to form an 

excipulum-like zone below the hypothecium. Pycnidia often present but inconspicuous, immersed, 

blackish, c. 80-200 /xm diam, ostiole often gaping; with a single ± globose locule but 

walls often convoluted with up to 5 locules seen in a vertical section; outer walls reddish brown, 

K-; inner walls of secondary locules hyaline; conidiogenous cells irregularly cylindrical, often 

with 1-2 percurrent proliferations, 7-10x1-1-5 /xm, base sometimes swollen to 2-7 /xm wide; 

conidia (mesoconidia) cylindrical, sometimes faintly biguttulate but not constricted in the 

middle, (3-8-)4-5(-5-5)x0-9-l-4/xm. 

Chemistry: All parts K-, C-, PD-; t.l.c: no substances detected. 

Observations: The apothecia of M. lutulata have a similar internal pigmentation to those of M. 

botryoides {q. v. ) and M. muhrii {q. v. ), but the species is most often confused with M. sylvicola. 

The latter shares the same habitat as M. lutulata, but can be distinguished by its hypothecium 

which always has a distinctly green, or rarely purplish, tinge in water mounts or KOH; in HNO3 

the pigment(s) turn purple-red. M. sylvicola further differs in its larger spores, taller hymenium, 

slightly longer and broader conidia, and presence of greenish pigment in its pycnidial walls. M. 

myriocarpa occurs in similar habitats to M. lutulata and has a reddish brown, but never opaque 

(in thin section) hypothecium. Furthermore, M. myriocarpa has smaller apothecia, narrower, 

often 1-septate spores, and sessile pycnidia with much shorter conidia.

150 


Map 11 Micarea lithinella ® 1950 onwards + Micarea lutulata • 1950 onwards O Before 1950 

Habitat and distribution: M. lutulata is found on rocks and sometimes exposed roots in dry 

rocky underhangs, and is a characteristic species of the Micareetum sylvicolae. It is apparently 

widespread in northern and western Britain; and although not yet reported from the south-west 

peninsula of England or the north-west of Scotland, it almost certainly occurs in those areas. It 

appears to be Uttle known outside of the British Isles, but I have seen material from Norway 

(Oppland and Hordaland), Sweden (Varmland), southern Finland, Germany (Baden- 

Wiirttemberg), and Czechoslovakia (Bohemia: Krkonose). 

Exsiccata: Arnold Lich. Exs. 409B p.p. (WRSL). Havaas Lich. Exs. Norv. 571 (EG). Larbal. Lich. 

Herb. 223 (BM, BM ex K). Rasanen Lich. Fenn. 612 p. p. (LD-mixed with M. tuberculata) 

22. Micarea melaena (Nyl.) Hedl. 

'(Figs21A,46A;Mapl2) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 82, 96 (1892). - Lecidea melaena Nyl. in Bot. Notiser 

1853: 182 (1853). - Lecidea vernalis var. melaena (Nyl.) Nyl. in Mem. Soc. Imp. Sci. nat. Cherbourg. 3: 

182 (1855). - Bacidia melaena (Nyl.) Zahlbr. in Annls mycol. 7: 474 (1909). Type: Sweden, on lignum, 

E. M. Fries, Lich. Suec. Exs. 212B (UPS-lectotype!; isolectotypes: C!, H-NYLp.m. 4778 [fragment]!, 

M!). 

Lecidea milliaria var. turfosa Fr. , Nov. Sched. Crit. 8: 7 (1826), non Biatora turfosa Massal. Type: Sweden, 

[? Smaland], on peaty turf, E. M. Fries, Lich. Suec. Exs. 212A (UPS-lectotype!; isolectotypes: C!, M!). 

.


Biatora stizenbergeri Hepp, Flecht. Eur. no. 504 (1860). Type: Switzerland, Rifferschwell, on dry plant 

debris amongst roots in peat moor, Hegetschweiler (BM - lectotype!; isotypes distributed in Hepp. 

Flecht. Eur. 504: BM!, E!, WRSL!). 

Lecidea ilyophora Stirton in Scott. Nat. 5: 220 (1879). Type: Scotland, Perthshire, Black Wood of 

substances detected]). 

Rannoch, on lignum, ix 1879, J. Stirton BM - lectotype! [t.l.c. : no 

Lecidea melaena f. catillarioides Vainio in Medd. Soc. Fauna Fl. fenn. 10: 12 (1883). - Micarea melaena f. 

catillarioides (Vainio) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 83, 96 (1892). Type: 

Finland, Karelia borealis, Nurmes, Riihivaara, on bryophytes on rock, 1875, Vainio (TUR-VAINIO 

21478 -holotype!). 

Lecidea melaena f. endocyanea Vainio in Medd. Soc. Fauna Fl. fenn. 10: 12 (1883). - Micarea melaena f. 

endocyanea (Vainio) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill 18 (3): 83, 96 (1892). Type: 

USSR, Karelia keretina, Kivakka, on mosses on rocks, 1877, Vainio (TUR-VAINIO 21476 - hole- 

type!). 

Bilimbia melaena f. aeruginosa Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 254 (1922). Type: Finland, 

Tavastia australis, Hollola, on lignum, 1875, Vainio (TUR-VAINIO 21461 - lectotype!). 

Bilimbia melaena f. epiphaeotera Vainio in Acta Soc. Fauna Fl. Fenn. 53 (1): 255 (1922). Type: Finland, 

Karelia borealis, Nurmes, Louhivaara, 'Kallion syrjalla', on bryophytes and old Cladonia squamules, 

1875, Vainio (TUR-VAINIO 21472- lectotype!). 

?Bilimbia melaena var. alnicola Savicz in Izv. imp. S. Peterb. bot. Sada 14 (I, Suppl.): 53 (1914). Type: 

USSR (?LE); request for loan received no reply. 

Bilimbia milliaria y. [var.] saprophila Korber, Parerg. lich. : 171 (1860); nom. superft. (Art. 63). 

Thallus effuse, superficial, of small, scattered to coherent or clustered, granular areolae 

(20-)25-80(-100) /xm diam, sometimes forming an uneven crust up to c. 150 fxm thick. Areolae 

very variable in colour, pale buff or pale dull green to dark grey-green; in section without an 

amorphous covering layer, outermost hyphae often with dark green (K-, HNO3-I- red) walls. 

Thallus often scurfy granular and blackish (especially in damp situations) owing to its disruption 

by invading dematiaceous fungi and foreign algae; in dry situations (e.g. undersides of fallen 

trunks) it may have a whitish or pale yellowish , farinose appearance due to a thick covering of an 

epiphytic alga, each cell of which is coated by hyaline, crystalline incrustations. Phycobiont 

micareoid, cells c. 4-7 jxm diam. 

Apothecia numerous, immarginate, convex-hemispherical to ± globose, sometimes tubercu- 

late, black, matt or slightly glossy, 0- 12-0-4 mm diam, or to 0-5 mm diam when tuberculate. 

Hymenium 30-40(-50) /xm tall, dull to bright aeruginose or grey-blue, K— or K-l- green 

intensifying; alternatively lower part (or rarely the entire hymenium) dilute purple-brown, K-l- 

greenish or K-l- purple intensifying; pigment(s) often more dense around the apices of the 

paraphyses, thus appearing as a dark 'epithecium' (c. 2-10 jxm tall) that contrasts with the more 

dilute colouration of the rest of the hymenium. Asci clavate, c. 30-40-12-15 jxm. Spores 

ovoid-oblong or oblong, with rounded apices, straight, (l-)3(-5)-septate, 12-21 x4-5(-5-5) /xm. 

Paraphyses numerous, branched and anastomosing, sometimes a few unbranched, thin, 0-8-1 

îm wide; apices not swollen, or sometimes slightly incrassate and up to 1-5 jxm wide, hyaline but 

often individually surrounded by a pigmented hood of dense gel (easily detached in mounts in K, 

by tapping the cover slip). Occasionally intermixed with ordinary paraphyses are a few which are 

simple , stout (c. 1 -5-2 jxm wide) and coated with pigment throughout their length so as to appear 

3-4 wide. Hypothecium c. 80-160 îm tall, dark and ± black in thick sections, pigmentation 

variable (see 'observations' below), usually purple-brown and K— or K-l- purple intensifying, 

or, outer (more rarely whole) part K-l- olivaceous with central part ('core') remaining ± purplish 

brown; all parts HNO3-I- purple-red; hyphae c. 0-7-1-5 jxm wide but their walls coated with 

dense pigment so that they appear to be c. 3-4 /xm wide, interwoven or becoming vertically 

orientated towards the hymenium, sometimes a few extending into the hymenium as stout 

pigmented paraphyses; ascogenous hyphae similarly pigment, short-celled, c. 2-5 /am wide. 

Excipulum indistinct, sometimes evident in sections of young apothecia, green or purplish 

(usually darker than hymenium, but paler than hypothecium); hyphae radiating, branched and 

anastomosing, c. 0-7-1 jxm wide. 

Pycnidia rare, black, of two types: (a) c. 100-140 /xm diam, sessile; walls purple-brown or dull 

olive, K-l- green; conidia {macroconidia) curved or hamate, 0-7-septate, 18-33 x 1-1-5 /xm; (b)


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

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

Chemistry: Thallus K— , C+ 

red or C— , PD; sections of apothecia C— ; t.l.c: gyrophoric acid 

(healthy thalli), or no substances or gyrophoric acid in trace amounts (poorly developed or 

disrupted thalH). 

Observations: Micarea melaena is one of the commonest species of the genus, especially in 

northern Europe, and it displays a considerable amount of variation in thallus appearance and 

internal apothecial pigmentation. However, my examinations of over 200 specimens have 

revealed many intermediates between the known extremes. When well-developed, the thallus is 

composed of cohering, small, granular areolae and is pale buff to dull green in colour. In 

exposed situations the thallus often becomes a dark green-black due to the more intense 

pigmentation of the outermost hyphae of the granules. The appearance of the thallus can also be 

affected by the presence of other organisms (see description and p. 00). When on lignum, in dry 

exposed situations the thallus may become much reduced to a fine scattering of tiny (less than 30 

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

during my explorations in Sweden. 

M. melaena also exhibits much variation with regard to the colouration of its apothecial tissues 

(especially the hypothecium). This is due to the presence of three (probably biosynthetically 

related) pigments, varying in their relative amounts and distribution. These pigments, all of 

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

intensifying; pigment B: purple, K+ green; pigment C: purple, K+ purple intensifying. 

In the epithecium and hymenium pigment A is often found alone, but sometimes it is replaced 

in the lower hymenium (or in extreme cases throughout the entire hymenium and epithecium) 

by pigment C. Pigment C is usually dominant in the hypothecium, but is often replaced in the 

upper and outer parts (or more rarely ± throughout) by pigment B. If thin sections of apothecia 

in which pigment B is dominant are mounted in K the lower central ('core') of the hypothecium 

retains a purplish tinge (the rest having turned green), indicating the presence of a small amount 

of pigment C. The same type of variation, presumably involving identical (or very similar) 

pigments, is found in several other species, e.g. M. crassipes and M. sylvicola. 

The pycnidia of M. melaena are very inconspicuous and apparently sparingly produced. 

However, a close scrutiny of about 100 specimens revealed ten collections with the microconi- 

dial state and only two (e.g. Coppins 6041) with the macroconidial state. A mesoconidial state 

was not found but may well occur. 

Habitat and distribution: M. melaena is widely distributed and especially common in northwest 

Europe. Although, for a Micarea, it is a conspicuous species, it is often overlooked as a 

black or charred stain on the tops of stumps and peaty turf! It is a common inhabitant of stumps 

and fallen decorticate trunks in upland woodlands, or in lowland woodlands on very acid, 

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

upland woods it can be found on the corticate trunks of healthy trees (especially Betula, 

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

conurbation) it has been found on the acidified bark of Acer and Ulmus. Occurrences on 

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

upland districts. Lichens associated with M. melaena on lignum and bark in the British Isles 

include Bryoria capillaris, B. fuscescens, Cladonia spp., Hypocenomyce scalaris, Hypogymnia 

physodes, Lecidea granulosa agg., L. icmalea, Micarea denigrata, M. prasina, Mycoblastus 

sterilis, Ochrolechia turneri, Parmelia saxatilis, Parmeliopsis spp., Platismatia glauca, Trapelia 

corticola, and Usnea spp. 

M. melaena is often terricolous, growing on oligotrophic, peaty soils, in montane regions or 

lowland heaths. In the English lowlands it is recorded from heaths in the Sussex Weald and east 

Norfolk. It is common on ± bare peat in the Pennines and Scottish highlands, but in the latter it 

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

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


-^y^ 

Map 12 Micarea melaena # 1950 onwards O Before 1950 

C. floerkeana, C. subcervicornis, Coelocaulon aculeatum s. lat., Lecidea icmalea, Micarea 

leprosula, and Ochrolechia androgyna. M. melaena is often found on rocks, growing over peaty 

debris or decaying mats of bryophytes and Cladonia squamules, but occurrences where it is 

growing directly on rock are rare. To my knowledge, all such occurrences in Britain are confined 

to woodland situations in rather heavily polluted areas approximating to zones 3-5 on the scale 

of Hawksworth & Rose (1970); these areas include Leicestershire, the West Yorkshire 

conurbation, and Fife. 

M. melaena is widely distributed in mainland Britain, but is particularly common in the upland 

areas of the north and west. There is a dearth of records from Ireland, where it is unlikely to be 

rare in suitable terrain. M. melaena is known from most parts of the European mainland and is 

particularly common in the boreal regions of Fennoscandia and upper Bavaria and adjoining 

alpine districts, but this apparent tendency may, at least partially, be a reflection of the avidity of 

the collectors active in those areas, and the herbaria which I have studied! It has been found in 

the Azores on Cryptomeria and sulphur-rich soils near fumaroles, but it is so far unrecorded 

from the Canary Islands. From outside Europe I have seen several collections from eastern 

Canada and north-eastern U.S.A. 

Exsiccata: Anzi Lich. Exs. Ital. 259A (BM, M). Anzi Lich. Sondr. 170B (UPS). Arnold Lich. Exs. 

332A-C (M). Arnold Lich. Mon. 49 (M), 248 (C, M), 249 (C, MANCH), 407 (C, M, MANCH). Britz. 

Lich. Exs. 946 (M). Fellman Lich. Am. 159 (H). Flotow Lich. Exs. 129C (UPS). Fries Lich. Suec. 212A, B 

(C, H-NYL p.m. 4778, M, UPS). Hepp Fl. Eur. 504 (BM, E, WRSL). Johnson Lich. Herb. 376 (BM). 

Krypt. Exs. Vindob. 362 (BM, M). Leighton Lich. Brit. 120 (BM). Malme Lich. Suec. 27 (C, M, S). Moug.


& Nestl. Stirp. Crypt. 1329 p.p. (E). Mudd Lich. Brit. 159 (M, MANCH). Norrlin & Nyl. Herb. Lich. 

Fenn. 180 (BM, C, M). Oliv. Lich. Orne 237 (M). Rasanen Lich. Fenn. 963 (M). Rasanen Lichenoth. 

Fenn. 344 (BM). Roum. Lich. Gall. 231 (M). Schaerer Lich. Helv. 196 p.p. (E). Vezda Lich. Set. 14 (BM, 

M). Zwackh Lich. Exs. 657, 675 (M). 

23. Micarea melaenida (Nyl.) Coppins, comb. nov. 

(Fig. 22) 

Lecidea melaenida Nyl. in Flora, Jena 48: 146 (1865). - Catillaria melaenida (Nyl.) Arnold in Flora, Jena 

53: 475 (1870). Type: France, [? Eure], Perrieres, Lenormand (H-NYL 18843 -holotype!). 

Lecidea relicta Nyl. in Flora, Jena 48: 354 (1865) [nee Stirton (1876)]. Type: France, Calvados, Falaise, 

L.-A. De Brebisson (H-NYL 21099 - holotype!). 

Catillaria schumannii {'SchumannV] Stein in Cohn, Krypt. -Fl. Schles. 2 (2): 232 (1879). - Lecidea 

schumanni Korber, Syst. Lich. Germ.: 255 (1855); nom. inval. (Art. 34). Type: East Germany, 

Reichenbach, near Ernsdorf, Schumann (WRSL - lectotype!; WRSL - isolectotype!). 

Catillaria zsakii Szat. in Magy. hot. Lap. 24: 108 (1926). - Toninia zsakii (Szat.) Lettau in Feddes Reprium 

Spec. nov. veg. SI: 9 (1955). Type: holotype not seen (? in BP); topotypes: Hungary, 'In argillosis 

natronatis prope oppidum Karcag, Comit. Jasz-Nagykun-Szolnok. Allit. ca. 80 m. s.m. Mens. Szept. 

1926', Z. Zsdk cfe G. Timko, Fl. Hung. Exs. 714 (E! [t.l.c: no substances], BM!, BM ex K!), and, from 

same locality but at '100 m. s.m.' and no date, Krypt. Exs. Vindob. 3154 (BM!, BM ex K!). 

Catillaria schumanni var. meridionalis Roux & Vezda in Vezda, Sched. Lich. Sel. Exs. 537 (1967). Type: 

France, Gard, Pujaut near Avignon, alt. 100 m, 'supra solum argillaceo-sabulosum, ad marginem 

Querceti ilicis', 6 xi 1966, C. Roux, Vezda Lich. Sel. Exs. 537 (BM - isotype!). 

Thallus effuse, terricolous (on argillaceous soils), forming a thin ± smooth, dull greenish 

white crust, c. 40-100 /xm thick, or forming confluent, convex areolae 0- 1-0-6 mm diam; crust 

sometimes becoming cracked so as to appear ± 'squamulose'. Thallus in section without a 

distinct hyaline, amorphous, covering layer, but with a hyaline 'cortex' 12-20 fxm tall, composed 

of compacted, evacuated hyphae (cytoplasm presumed lost owing to lack of staining by LCB or 

ammoniacal erythrosin); thallus hyphae l-5-2-5(-3) fxm wide; numerous algae-free hyphae 

penetrating the soil substratum to a depth of c. 1 mm (? or more). Phycobiont micareoid, cells 

4-7 /Lim diam. 

Apothecia numerous, scattered or 2-4-confluent, immarginate, convex to hemispherical, 

black, matt, 0-2-0-6 mm diam; old, contorted apothecia sometimes reaching 1-2 mm diam; 

occasionally a fine weft of radiating, white (hyaline) hyphae is seen around the edge of an 

apothecium. Hymenium 35-40 ^tm tall; uppermost c. 1-5 /am (epithecium) with dense, 

purple-red; remaining (lower) parts dilute 

dull purple, K— , HNO3+ red. Asci clavate, 35-38x9-12 /xm. Spores ellipsoid, oblong-ellipsoid 

or ovoid-oblong, straightor slightly curved, 0-1-septate, (7-)9-5-15x3^(-4-5) /am. Paraphyses 

numerous, mostly simple below, but often branched above, sometimes anastomosing, 1-1-7 ixm 

wide; apices (upper 5-15 /xm) often purple-brown pigmented and to 2-5 /xm wide. Hypothecium 

amorphous, purple-brown pigment, K— , HNO3+ 

60-120(-200) /Ltm tall, mottled dark purplish brown, K— , or K+ purple intensifying in part, 

HNO3-I- purple-red; hyphae interwoven but ± vertically orientated in uppermost part, c. 1-1-8 

/am wide, each with a densely pigmented gel sheath thereby giving a total diameter of c. 3 /xm; 

ascogenous hyphae c. 2-5-5 /am wide. Excipulum soon reflexed, but distinct in sections of young 

apothecia, purple-brown (usually darker than hymenium); hyphae radiating, branched and 

anastomosing, c. 1-3-2 /xm wide, embedded in densely pigmented gel-matrix. 

Pycnidia often present, immersed, black, c. 50-100 /xm diam; walls purple-brown, K— or Kf-Ipurple 

intensifying; conidia (? microconidia) , ± cylindrical, 4-8-6(-7)xl /xm. 

Chemistry: Thallus K— , C— , KC— 

, PD-; t.l.c: no substances. 

Observations: Micarea melaenida is characterized by its whitish, terricolous thallus, immar- 

ginate, black apothecia (with purplish pigmentation in section), numerous and rather stout 

paraphyses, and 0-1-septate spores. It is apparently related to M. assimilata {q. v. ) and its allies. 

When compared with M. melaenida, forms of M. melaena with immature spores can be 

distinguished by their finely granular areolae, narrow and less numerous paraphyses, and usual


presence of green pigments in apothecial sections (in water or K mounts). M. turfosa is similar in 

some ways to M. melaenida, but has a dark coloured thallus, an olivaceous hymenium, reddish 

brown (never purplish) hypothecium, and larger spores which are often 1-3-septate. 

Habitat and distribution: M. melaenida grows on, and in direct contact with, fine-grained 

sandy or argillaceous, mineral soils, and is not known to occur on decaying bryophytes or plant 

debris, etc. Unfortunately I have never seen M. melaenida in the field, and there is little 

ecological information to be gleaned from the literature or herbarium labels. Furthermore, the 

specimens examined have included few associated species, and only Cladonia sp. (squamules) 

and Catapyrenium lachneum have been noted. In its Hungarian locality it was said to have 

occurred on soil rich in precipitated salts ('argillosis natronalis') although no effervescence was 

apparent when I tested the soil with 50% HNO3. It is probable that M. melaenida is restricted to 

niches where the salts have been permanently leached out. A sample of soil from Fl. Hung. Exs. 

714 was sent for analysis to the Macaulay Institute for Soil Research (Aberdeen) and Dr B. W. 

Bache of the Institute reports 'It bears no relationship at all to a sodic soil, and is in fact fairly 

acid, with pH 5-2. The conductivity is fairly low (0-8 mS in 1:2 extract), so it does not contain any 

great concentration of soluble salts, nor does it contain any carbonate.' 

The data on the herbarium labels, although limited, do indicate that it occurs at low altitudes, 

for example the collection from Loire Atlantique was made from soil on coastal cliffs; that from 

Gard was made at 100 m; and those from Hungary were made at 80-100 m. 

M. melaenida is a rare species with a rather southern distribution, and I have seen material 

from several localities in France (e.g. Loire Atlantique, Vienne, and Gard), and single localities 

in East Germany (Dresden) and Hungary. It is not known from Scandinavia, and its status as a 

British plant is doubtful. Its first British report seems to be its appearance in the checklist of 

James (1965fl), but I have been unable to ascertain the reason for its inclusion. However, its 

presence in north-west France suggests that it could occur in Britain, especially along the south 

coast of England or in the Channel Islands. 

Material from South Africa and distributed as Almborn Lich. Africani no. 84 is not M. 

melaenida but a mixture of two, undescribed species, both of which are probably referable to 

Micarea. 

Exsiccata: Fl. Hung. Exs. 714 (BM, BM ex K, E). Krypt. Exs. Vindob. 3154 (BM, BM ex K). Vezda 

Lich. Sel. 537 (BM). 

24. Micarea melaenizaHedl. 

(Figs21B,38B) 

in Bih. K. svenska VetenskAkad. Hand. Ill, 18 (3): 83, 96 (1892). - Lecidea melaeniza ["melaniza'] 

(Hedl.) Zahlbr., Cat. lich. univ. 3: 798 (1925). Type: Sweden, Halsingland, lignum of old pine trunk, viii 

1891,7. T. Hedlund iS-ho\otype\). 

Thallus effuse, lignicolous (sometimes extending on to bryophyte or lichen thalU), endoxyhc, 

or epixylic with whitish convex areolae; areolae c. 100-400 fxm diam and up to 45 /am thick, 

without an amorphous covering layer. Phycobiont micareoid, cells 4-7 /xm diam. 

Apothecia numerous, few, or absent, immarginate, at first convex-hemispherical or subglobose, 

often becoming tuberculate, black, matt, 0-2-0-3 mm diam, or to 0-5 mm diam when 

tuberculate. Hymenium 28-42 /xm tall, upper and lower parts aeruginose-green, middle part ± 

hyaline with aeruginose vertical streaks; K- or dulling, HNO3-I- red. A^dclavate, 26-35 x 10-12 

yarn. Spores ovoid or dacryoid, simple, 5-9x2- 5-3 -8 yam. Paraphyses rather numerous, of two 

types; p.p. evenly distributed, flexuose, branched, thin, c. 0-7-1 ^tm wide, hyahne, but apices 

sometimes pigmented and then to 2-5 /am wide; p.p. scattered or in small fascicles, stout, c. 2 /am 

wide, each coated ± throughout their length by greenish pigment. Hypothecium c. 120-200 /am 

tall, dark brown, sometimes with a faint reddish (never purplish) tinge, K- or dulling, HNO3 - 

or red tinge slightly intensifying; hyphae coated with dark brown pigment and c. 2-3 /am wide, 

interwoven but becoming vertically orientated towards the hymenium and sometimes con-


tinuing into it as stout pigmented paraphyses; ascogenous hyphae similarly pigmented, but with 

short swollen cells to 5 /xm wide. Excipulum not evident in available material. 

Pycnidia numerous, black, sessile and c. 40-60 /xm diam, or more usually borne on black 

stalks (pycnidiophores) and 80-300x40-70 îm (including stalk); stalks simple, or sometimes 

branched and bearing up to four pycnidia; upper part (pycnidial wall) dark olive but lower part 

(stalk tissue) reddish brown, both parts K— , Conidia {mesoconidia) short cylindrical, 2-3- 

3-6xl-l-3/>im 

Chemistry: Thallus PD— ; sections of apothecia and thallus C-; material insufficient for 

analysis by t.l.c. 

Observations: With its black, subglobose to tuberculate apothecia, black, stalked pycnidia, 

inconspicuous thallus, and occurrence on lignum, M. melaeniza is indistinguishable in external 

appearance from M. misella and M. nigella. Microscopically, M. misella can be distinguished by 

the olivaceous (K+ violet) pigment in its hymenium and pycnidia, and its ± hyaline hypothecium; 

and M. nigra can be distinguished by the purple-brown (K+ green, HNO3+ purple-red) 

pigment in its hymenium, hypothecium and pycnidia. With respect to apothecial and pycnidial 

pigmentation and structure, M. melaeniza is almost identical to M. botryoides. However, M. 

botryoides is rarely lignicolous, and has larger, often septate spores, and longer mesoconidia. M. 

muhrii has similar apothecial pigments to those of M. melaeniza, but is readily distinguished by 

its larger, adnate apothecia (which often have a pale marginal rim), larger spores, well 

developed excipulum, and lack of stalked pycnidia. 

Habitat and distribution: M. melaeniza is a rare species, known only from three collections 

made in Sweden (Angermanland, Halsingland, and Smaland) where it occurred on the lignum 

of conifer trunks. Associated species identified on the examined collections include Calicium 

salicinum, Cetraria pinastri, Chaenothecopsis lignicola, Cladonia spp. (squamules), Dimerella 

diluta, Hypogymnia physodes, Lecanora symmicta agg., Micarea anterior, M. misella, M. 

prasina, Usnea sp., and Lophozia sp. 

25. Micarea melanobola (Nyl.) Coppins, comb. nov. 

(Figs 21c, 46B-C) 

Lecidea melanobola Nyl. in Flora, Jena 50: 371 (1867). - Catillaria melanobola (Nyl.) Vainio in Acta Soc. 

Fauna Fl. fenn. 57 (2): 465 (1934). - Lecidea erysiboides *L. melanobola (Nyl.) Nyl. in Hue in Revue Bot. 

Courrensan 5: 103 (1886). - Catillaria |3. byssacea f. melanobola (Nyl.) Blomb. & Forss., Enum. PI. 

Scand. : 91 (1880). - Micarea prasina f . melanobola Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 

(3): 87 (1892). Type: Finland, Tavastia australis, Kuhmois [Kuhomoinen], on bark of Picea abies , 

/. P. Norrlin (H-NYL 21614 -lectotype!; isolectotypes: H-NYLp.m. 4504!, H!). 

Thallus effuse, thin and indistinct, of scattered to ± coherent, minute olivaceous granules 

(goniocysts), c. 12-25 jxm diam, which appear ± gelatinous when moist. Outermost hyphae of 

goniocysts with greenish, K-l- violet walls. Phycobiont micareoid, cells 4-7 (xm diam. 

Apothecia numerous, convex-hemispherical, immarginate, dark grey to black, 0- 1-0-24 mm 

diam. Hymenium 30-35 /xm tall, hyaline, but with a clearly delimited, dark green epithecium, 

K+ deep violet, C+ violet, HNOs-l- red; pigment closely bound to (? or also in) the apical walls 

of the paraphyses, and also present in the gel-matrix and apical walls of the asci. Asci clavate, 

30-35x9-11 fjLm, upper wall(s) sometimes tinged with greenish, K+ violet pigment. Spores 

ellipsoid, ovoid, or oblong-ovoid, with obtuse apices, 0-1-septate, 7-9-7x2-5-3-3 /xm. Paraphyses 

scanty, branched and anastomosing, thin, 0-5-1 /xm wide; apices thickened with greenish 

(K-l- violet) pigment and up to 1-7 /xm wide, often overtopping the tops of the asci. Hypothecium 

25-70 /xm tall, hyaline or dilute dull yellowish. Excipulum indistinct, sometimes evident as a 

non-amyloid, narrow (c. 10-15 /xm), reflexed zone of radiating, branched and anastomosing 

hyphae, 0-5-1 /xm wide. 

Pycnidia numerous but small and inconspicuous, sessile, black, with dark greenish (K-l- 

violet) walls; of two types: (a) 40-50 /xm diam; conidia (mesoconidia) cylindrical or ovoid- 

1866, 

I


Table 6 Diagnostic features for the separation of Micarea melanobola, M. misella, and M. prasina.


26. Micarea misella (Nyl.) Hedl. 

(Fig. 23A; Map 13) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 78, 88 (1892). - Lecidea anomala f. misella Nyl., 

202 (1861) .- Lecidea misella (Nyl.) Nyl. in Not. Sdllsk. Fauna Fl.fenn. Fork. 8: 177 (1866). 

Lich. Scand. : 

Type: Finland, Nylandia, Helsingfors [Helsinki], 1858, W. Nylander (H-\ectotypel). 

Lecidea resinae* [subsp.] L. globularis Nyl. , Lich. Scand. : 213 (1861). - Lecidea globularis (Nyl.) Lamy in 

Bull. Soc. bot. Fr. 25: 435 (1878). - Micarea globularis (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. 

Handl. Ill, 18 (3): 78, 88 (1892). Type: Sweden, 'Arthonia turgida var. y globifera Svecia 112°' (H-ACH 

52-holotype!). 

Lecidea melanochroza Leighton ex Crombie in 7. Bot., Lond. 9: 178 (1871). Type: Scotland, Perthshire, 

near Loch Tummel, /. M. Crombie (BM - lectotype!; isolectotypes: BM ex K!, and distributed in 

Crombie Lich. Brit. Exs. 174 (BM!) [note: this example includes M. prasina and those in E and M 

contain M. prasina only]). 

Lecidea misella f. brasiliana Vainio in Acta Soc. Fauna Fl. fenn 7 (2): 57 (1890). Type: Brazil, Minas 

Geraes, Carassa, 1400 m, on lignum, Vainio, Lich. Brasil. Exs. 1420 (BM ex K - lectotype!; M - 

isolectotype!). 

Bilimbia melaena f. decrustata Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 255 (1922). Type: Finland, 

Tavastia australis, HoUola, 1871, Vainio (TUR-VAINIO 21480- lectotype!). 

Lecidea asserculorum sensu Th. Fr., Lich. Scand. 2: 473 (1874), non Ach (1810). See note by Hedlund 

(1892: 89). 

Thallus effuse, often wide-spreading, usually lignicolous and endoxylic, sometimes develop- 

ing on the surface of the substratum as convex to irregularly subglobose, greenish white to 

greenish grey, ± contiguous granular-areolae, c. 60-120(-150) /w-m diam. Areolae in section, 

without a well defined cortex or hyaline amorphous covering layer, but outer 5-12 /xm is often 

tinged with olivaceous-brown pigment (K+ violet); pigment confined to gel-matrix and not 

adhering to hyphal walls. Phycobiont micareoid, cells 4—7 /xm diam. 

Apothecia numerous, few or absent, scattered to crowded and confluent, immarginate, 

convex-hemispherical to subglobose, sometimes shortly stipitate, sometimes tuberculate, black, 

rarely pallid or grey-brown (shade forms), matt. c. 0-1-0-3 mm diam. Hymenium 25-36 /am tall; 

upper part dull greenish brown or olivaceous, K-l- violet, with pigment confined to the 

gel-matrix; remaining (lower) part hyaline or dilute sordid greenish. Asci clavate, 25-35x7-10 

^tm. Spores ellipsoid, ovoid, or oblong-ovoid, not curved, sometimes with 1-3 guttules, simple 

but a few (very rarely many) sometimes 1-septate, (6-5-)7-9-5x2-0-3(-3-7) jxm. Paraphyses 

scanty, sparingly branched 0-5-0-8 /xm wide; apical parts often more richly branched, sometimes 

wider (to 1-5 /xm), but without closely adhering pigment. Hypothecium c. 45-70 /xm tall, 

hyaline, or dilute olivaceous (rarely dark olivaceous: 'f. brasiliana') and then K-f- violet; hyphae 

interwoven but becoming vertically orientated towards the hymenium, c. 0-7-1 /xm wide; 

ascogenous hyphae with swollen cells, c. 2-4 /xm wide. Excipulum poorly developed but often 

evident as a dark brownish (K+ violet) zone bordering the reflexed edges of the hymenium; 

hyphae radiating, branched and anastomosing, very narrow, c. 0-5-0-8 /xm wide. 

Pycnidia usually present black, walls green-brown, K-l- violet; of two types: (a) sometimes 

sessile, more usually borne single in the top of a black, unbranched stalk (pycnidiophore), 

70-320 /xm tall (including stalk) and 50-100 /xm wide; stalk part composed of interwoven 

hyphae, c. 1-1 -5 /xm wide, embedded in a gel-matrix containing a dull brown or olivaceous, K-lviolet 

pigment; conidia (mesoconidia) shortly cylindrical, sometimes faintly biguttulate, 3-5- 

5xl-l-5(-l-7) /xm; (b) ± immersed in the substratum (lignum) or areolae, c. 40 /xm diam; wall 

with green-brown, K-l- violet pigment in upper part, but becoming ± hyaline below; conidia 

(microconidia) narrowly cylindrical, 3-8-6x0-6--0-8 /xm. Pycnidia with stalks (a) are mostly 

confined to forms with an endoxylic thallus. 

Chemistry: Sections of thallus (when superficial) K- , C-l- red, PD- ; sections of apothecia C-fviolet 

(olivaceous pigment), but only rarely C-l- red; t.l.c: gyrophoric acid detected in varying 

amounts, sometimes absent. 

Observations: Typical forms of M. misella with an endoxylic thallus, numerous apothecia, and 

stalked pycnidia are easy to recognize. However, when it has a well developed superficial thallus

LICHEN GENUS MICAREA IN EUROPE 

Table 7 Diagnostic features for the separation of Micarea denigrata and M. misella. 

Apothecia size (mm) 

Thallus type 

Spore septation 

Spore length (/u.m) 

Paraphyses 

Hymenium 

Macroconidia (/u,m) 

Mesoconidia (/Am) 

Microconidia (/u,m) 

Pycnidia (meso-) 

Chemistry (t.l.c.) 

0-1-0-3 

Usually endoxylic; sometimes 

with areolae, 60-120 /x,m diam 

0(-l) 

6-5-9-5 

sparse; 0-5-0-8 )u,m wide 

C+ violet; very rarely also 

C+ orange-red 

unknown 

3-5-5 X 1-1-5 

3-8-5x0-6-0-8 

sessile to stalked 

nil; but gyrophoric acid 

sometimes present in trace 

amounts, rarely in large amounts 

misella denigrata 

0-1-0-5 

usually with areolae, 60-200 /nm 

diam; rarely endoxylic 

(0-)l 

(7-)9-16(-18) 

numerous; 1-1-7 jxm wide 

C+ violet; usually also 

C-l- orange-red 

12-24x1; curved 

2-8-4-5(-5)x 1-2-1-8 

(4-5-)5-7-5x 0-7-0-8 

immersed to emergent, never 

stalked 

gyrophoric acid usually present 

in large amounts; very rarely 

not detectable 

and no stalked pycnidia it can be difficult to separate from M. denigrata, especially if in the latter 

no conidial (or only the microconidial) state can be found. In such instances M. misella can 

usually be distinguished by its small apothecia, mainly simple, uncurved ascospores and sparse, 

thin paraphyses. The same criteria can separate forms of both species with an endoxylic thallus 

and lacking diagnostic macroconidial or mesoconidial states. A C+ orange-red reaction can 

usually be obtained in sections of the apothecia of M. denigrata, although the reaction is 

sometimes very weak or unobtainable in diminutive, lignicolous, epixylic forms; in M. misella I 

have obtained this reaction in only one or two extremely well-developed specimens. Table 7 

outlines the main diagnostic features of these two species. Apart from M. misella, black, stalked 

pycnidia are also a characteristic feature of M. botryoides, M. melaeniza, and M. nigella; 

however, the pycnidial wall tissue in these species is brownish or greenish in KOH, and never 

violaceous as in M. misella. 

For differences from M. melanobola and M. prasina see Table 6. 

Habitat and distribution: M. misella is almost exclusively lignicolous (especially on conifer 

lignum). It has not been found on bark, but has been collected on Picea cones, old basidiomes of 

polypores (Daedaleopsis confragosa and Gloeophyllum sepiarum), and on moribund Polytrichum 

spp. by woodland tracks. Most British collections of M. misella have been made from 

the sides and undersides of fallen decorticated trunks of Pinus, and associated species included 

Lecidea icmalea, Micarea prasina, Xylographa abietina, X. vitiligo, Cladonia spp. (scattered 

squamules), and Hypogymnia physodes. Reports on exposed roots and stones (e.g. Watson, 

1930: 53) refer to other species, especially M. bauschiana. In Britain M. misella appears to be 

rather rare and restricted to Scotland where it is mainly found in the native pine-woods. It has 

been reported (as 'Lecidea asserculorum') from England and Wales, but all the pertinent 

material seen belongs to diminutive forms of M. denigrata and M. prasina, or quite different 

species such as M. melaena and Cliostomum grifftthii. 

M. misella is widely distributed in Europe, especially in areas with naturally occurring 

coniferous forests. It is found as far north as Lycksele Lappmark and Norbotten in Sweden, and 

an outlying southerly locality is on the Mediterranean island of Corsica where it was collected on 

the rotting lignum of Eucalyptus. From outside Europe I have seen material of M. misella from 

Canada (Ontario and Newfoundland), and from the highlands of Minas Geraes (at 1400 m) in 

Brazil. 

Exsiccata: Arnold Lich. Exs. 626, 627 (BM ex K, M). Arnold Lich. Mon. Ill, 241, 307 (BM ex K, M, 

MANCH). Britz. Lich. Exs. 208 (M). Claudel «& Harm. Lich. Gall. 445 (BM). Crombie Lich. Brit. 11 

A 

159


Map 13 Micarea misella # 1950 onwards O Before 1950 

p.p. (BM). Elenkin Lich. Ross. 189 (UPS). Krypt. Exs. Vindob. 1532 (BM, BM ex K, M), 4214 (BM ex K, 

M). Malme Lich. Suec. 365 (S, WIS). Norrlin & Nyl. Herb. Lich. Fenn. 744 (H). Reliq. Suza. 42 (BM, 

GZU, WIS). Vainio Lich. Bras. 1420 (BM ex K, M), 1451 (BM). Zwackh Lich. Exs. 1085 (M). 

27. Micarea muhrii Coppins, sp. nov. 

(Fig. 23B) 

Thallus effusus, endoxylicus et labem albidam efficiens, vel epixylicus aut epilithicus; thallus superficialis 

cum areolis convexis, albidus, c. 0-06-0.2 mm diam, interdum coalescentibus in crustam crassiusculam 

demum rimosam. Algae cellulis 4-7 yam diam. Apothecia adnata, convexa vel hemisphaerica autem baud 

globosa, griseo-atra, aut pallida vel rufo-brunneola vel fumosa ubi umbrose, 0- 15-0-46 mm diam, 

immarginata sed vulgo zona marginali, complana, ± incolorata. Hymenium 40-45 jxxn altum, parte summa 

obscure olivacea vel griseo-viridi et K— , vel raro hyalina. Ascosporae ellipsoideae, ovoideo-ellipsoideae 

vel oblongo-ellipsoideae, simplices, 9-12x4-5 /u,m. Paraphyses aliquantum paucae, ramosae et interdum 

anastomosantes, 1-1-5 /u,m latae, apices versus vulgo incrasstae ad 3-5 /xm latae et vulgo pigmentiferae. 

hyalinum, paulum evolutum mox reflexum. 

Hypothecium obscure rufofuscum, K— , HNO3— 

. Excipulum 

Pycnidia pauca, immersa. Conidia cylindrica, 4-6x0-8-1 /x.m. Thallus et apothecia K-, C-, PD — ; sine 

materia chemica. 

Typus: Suecia, Wermelandia, par. Lungsund, Pungbacken, alt. c. 170 m, in loco aperto ad truncum 

decorticatum supra rivum, 15 vii 1980, leg. L.-E. Muhr 2851 (E-holotypus). [t.l.c: no substances.]


Thallus effuse, endoxylic and visible as a white stain, or epixylic, or rarely epilithic; superficial 

thallus formed of whitish, greenish-white or grey-white, convex areolae, c. 0-06-0-2 mm diam, 

which are especially well developed around the apothecia; areolae sometimes coalescing to form 

a thickish crust up to 0-2 mm thick, which may eventually become rimose. Areolae in section 

without a hyaline amorphous covering layer. Phycobiont micareoid, cells 4-7 /u,m diam. 

Apothecia numerous, occasionally coalescing, adnate, convex, soon becoming hemispherical, 

but never globose or tuberculate, grey-black, or pallid, reddish brown or brown-grey in shade 

forms, 0- 15-0-46 mm diam; disc matt or slightly glossy; immarginate but the adnate rim is often 

seen as a ± colourless or watery grey zone, c. 20-40 /xm wide. Hymenium 40-45 /xm, ± hyaUne 

with upper part dark olivaceous or grey-green, K— , HNOs-f- purple-red, or ± hyaline 

throughout in shade forms. Asci cylindrical-clavate, 38-40x10-12 /xm. Spores simple, often 

biguttulate, ellipsoid, ovoid-ellipsoid or oblong-ellipsoid, 9-12x4-5 /xm. Paraphyses rather 

scanty, branched (especially in their upper parts), sometimes anastomosing, 1-1-5 /zm wide, 

often sUghtly incrassate at apices to 2 /^m wide; or upper 9-15 /xm of paraphyses thickened by 

dense greenish pigment and then to 3-5 /xm wide. Hypothecium c. 70-120 îm tall, dark reddish 

brown, K— , HNO3— 

; hyphae c. 1-2 /xm wide embedded in the densely pigmented matrix, 

interwoven but becoming vertically orientated towards the hymenium; ascogenous hyphae with 

short, swollen cells to 5 (xm wide. Excipulum sometimes evident in sections of young apothecia 

as a non-amyloid, hyaline zone, but soon reflexed and obscured in older apothecia; composed of 

radiating, branched and anastomosing hyphae l-l-5(-2) îm wide. 

Pycnidia rare, sunken within the areolae, c. 40-50 fxm diam; upper part of wall around the 

ostiole olivaceous, lower part reddish brown, all parts K-. Conidia {microconidia) cylindrical, 

4-6x0-8-1 /xm. 

Chemistry: Thallus K— , C— , PD- ; apothecia sections C— ; no substances detected by t.l.c. 

Observations: Micarea muhrii is characterized by its convex-adnate, never tuberculate, 

usually dark grey apothecia, greenish upper hymenium, dark reddish brown hypothecium 

(without green or purple tinges, even in K), and simple spores. It recalls the mainly saxicolous 

M. lutulata in its pigmentation, but that species differs in having convex-subglobose, often 

tuberculate apothecia, smaller spores, and a larger celled phycobiont. The lignicolous M. 

melaeniza is another species with similar pigmentation, but has smaller, markedly convex to 

tuberculate apothecia, smaller spores, and sessile or stalked pycnidia. 

Habitat and distribution: Often in abundance on lignum of decorticate logs that lie across 

streams and probably become inundated at times; collected once on a periodically inundated 

boulder in a stream. So far, known only from Varmland in Sweden, from where it has been 

found in at least four localities by Lars-Erik Muhr, in whose honour I have the pleasure of 

naming this species. 

28. Micarea myriocarpa V. Wirth & Vezda ex Coppins, sp. nov. 

(Figs 23C, 47C; Map 14) 

Micarea myriocarpa V. Wirth & Vezda in Poelt & Vezda, Bestimmungsschl. europ. Flechten, Erganzungsheft 

I: 161 (1977); nom. nudum (Art. 32). - Micarea myriocarpa V. Wirth & Vezda in V. Wirth, 

Flechtenfl.: 341, 345 (1980); nom. nudum (Art. 32). 

Thallus effusus, farinoso-granulosus, tenuissimus vel crassiusculus ad 0-3 mm crassus, pallide viridis vel 

viridulo-fulvus. Algae cellulis ± globisis, c. 4-1 ^tm diam, conglomeratis in granula goniocystiformes. 

Apothecia immarginata, primum convexo-hemisphaerica mox tuberculata, pallide vel obscure rufa, vel 

fusca, 0-1-0-25 mm diam. Hymenium 25-35 /xm altum, ± hyalinum vel dilute rufo-brunneolum, cum vittis 

verticalibus obscure rufo-brunneolis, K — . Ascosporae oblongae vel oblongo-ovoideae, rectae vel ± 

curvatae, 5-5-8-5 x 1 •5-2-5 /xm. Paraphyses aliquantum paucae, dimorphae:p.p. hyalinae, laxae, simplices 

vel parce ramosae, interdum anastomosantes, graciles, 0-8-1-2 fxm latae, apicibus interdum incrassatis ad 

1-8 fim \ahs; p.p. pigmentiferae, fasciculatae, plerumque simplices, crassae, 2-3 /xm latae. Hypothecium 

rufo-fuscum vel armeniaco-fuscum, K — , HNO3 

- . Excipulum nullum. Pycnidia pauca et inconspicuosa, ±


sessilia, doliiformia, rufo-fusca, c. 25-30 /tm diam. Conidia breviter cylindrica, 2-5-3-2X 1-1-3 /u.m. Thallus 

et apothecia K— , C— 

, PD 

— ; sine materia chemica. 

Typus: Germania: 'Baden, Nordschwarzwald, Schurmsee bei Schonmiinzach bei Schwarzenberg, alt. 

790 m, Tannen am Seeausfluss', 25 v 1976, leg. V. Wirth (hb V. Wirth 6085 - holotypus!). 

Thallus effuse, pale green or greenish buff, scurfy farinose-granular, thin or thickish up to 0-3 

mm thick. Phycobiont ?micareoid, grouped into small goniocyst-like clusters c. 10-15 ^tm diam; 

cells ± globose 4-7 ixxn diam, or ellipsoid 5-7x3-5-5 /xm. 

Apothecia numerous, immarginate, at first convex-hemispherical, soon becoming ± globose, 

often becoming tuberculate, pale to dark reddish brown, rarely brown-black, 0- 1-0- 15 mm 

diam, or to 0-25 mm diam when tuberculate. Hymenium c. 25-35 ^x.m tall, hyaline or dilute 

brownish, with dark brown, vertical streaks due to fasciculate pigmented paraphyses; K-, 

HNO3— . Asci cylindrical-clavate, 21-25x5-7 jxm. Spores oblong or oblong-ovoid, straight or 

slightly curved, simple or 1 -septate, 5 •5-8-5x1 •5-2-5 ^cm. Paraphyses rather scanty, of two 

types: p.p. hyaline throughout, evenly distributed, simple or sparingly branched (especially 

above), sometimes anastomosing, thin, 0^8-l-2 /otm wide, sometimes widening towards their 

apices to 1-8 /xm; p.p. pigmented ± throughout, in small fasciculate clusters arising from the 

hypothecium, stout, 2-3 /am wide. Hypothecium 35-85 jxm tall, reddish brown or dull orange- 

brown, K-, HNO3— ; hyphae coated with dense brownish pigment and c. 2-3 /xm wide, 

interwoven but becoming vertically orientated towards the hymenium and sometimes continuing 

into it as pigmented, fasciculate paraphyses; ascogenous hyphae similarly pigmented but 

with short swollen cells, to 4 /xm wide. Excipulum not evident, even in sections of young 

apothecia. 

Map 14 Micarea myriocarpa # 1950 onwards O Before 1950


Pycnidia occasionally present but very inconspicuous, ± sessile, doliiform reddish brown, c. 

25-35 ixm tall and 25-30 /xm diam; wall reddish brown or dull orange-brown, K— , HNO3— 

Conidiogenous cells ± cylindrical to ampulliform, 4-5 x 1-1 -2 /xm but base sometimes swollen to 

2 jxm wide. Conidia (mesoconidia) short cylindrical, sometimes faintly biguttulate, 2-5-3-2xl- 

1-3 fxm. 

Chemistry: Thallus K— , C— , PD— ; apothecia sections C— ; no substances detected by t.l.c. 

Observations: Micarea myriocarpa is characterized by its numerous, small, ± globose to 

tuberculate, reddish brown apothecia, darkish (but never blackish, even in thick sections) 

hypothecium, and small, narrow, 0-1-septate spores. The apothecial tissues completely lack any 

greenish pigmentation, unlike in M. tuberculata and Psilolechia clavulifera, species of similar 

habitats with which M. myriocarpa has been confused. M. botryoides is another common 

associate of M. myriocarpa, but can be distinguished by its black apothecia, darker hypothecium, 

larger spores, and numerous stalked, black pycnidia. 

Habitat and distribution: M. myriocarpa is faithful to the Micareetum sylvicolae and is found 

especially on exposed roots in dry underhangs under trees on steep slopes in valley woodlands. 

In the same situations it has been collected also on loose stones and mats of dry bryophytes. An 

additional habitat for M. myriocarpa is the dry undersides of stumps (especially conifers), 

situations which are microclimatically identical to rocky underhangs. In Britain, M. myriocarpa 

is widely distributed, but little collected, in western and northern districts. On the Continent the 

species is little known, but I have seen material from western Norway as far north as Trondheim, 

and Varmland in Sweden; and Wirth {loc. cit.) records it from the northern Schwarzwald in 

Germany (the type locality). 

29. Micarea nigella Coppins, sp. nov. 

(Figs24A,47B;Mapl5) 

Thallus effusus, immersus vel tenuissimus, albidus vel viridi-griseus. Algae cellulis 4-7 /xm diam. 

Apothecia primum subglobosa mox tuberculata, immarginata, atra, 0- 1-0-3 mm diam. Hymenium 25-30 

/Ltm altum, ± incoloratum, autem parte summa (epithecio) purpureo-fusca K-l- olivacea. Ascosporae 

ellipsoideae, ovoideae vel oblongo-ovoideae, simplices 6-5-11 x2-5-4 /xm. Paraphyses aliquantum 

paucae, dimorphae: p.p. hyalinae, laxae, ramosae, gracillimae, 0-7-1 /u,m latae, apicibus vulgo leviter 

incrassatis ad 2 /xm latis; p.p. pigmentiferae, fasciculatae, plerumque simplices, crassae, 2-2-5 ^tm latae. 

Hypothecium obscure purpureo-fuscum, K+ obscure olivaceum. Excipulum paulum evolutum mox 

reflexum. Pycnidia numerosa, atra, sessilia vel stipitata, interdum ramosa, c. 40-300 )u,m alta et 40-80 /xm 

lata, parietibus purpureo-fuscis, K-l- obscure olivaceis. Conidia ellipsoidea ad breviter cylindrica, 3-4- 

4-3x1-2-1-6 ixm. Thallus et apothecia K-, C-, PD-. 

Typus: Dania, Jyllandia, c. 16 km septentriones e Hobro, Rold Skov, Torstedlund Skov, ad truncum 

decorticatum vetustum coniferarum in sylva mixta, 8 viii 1979, B. J. Coppins 4429 (E - holotypus). 

Thallus effuse, endoxylic, or ± epixyhc as a thin whitish to pale green-grey crust, but not 

forming discrete granules or areolae. Phycobiont micareoid, cells 4-7 pm diam. 

Apothecia few to numerous, immarginate, subglobose at first, often becoming tuberculate, 

black, matt, 0-1-0-2 pm diam, or to 0-3 mm diam when tuberculate. Hymenium 25-30 îm tall, 

hyaline or tinged dull purplish brown in places, often with darker, purplish brown vertical 

streaks; upper part (2-15 pm) irregularly pigmented purplish brown; K-h dull green, HNO3-Ired. 

Asci clavate, 22-27x10-11 pm. Spores ellipsoid, ovoid or oblong-ovoid, simple, 6-5- 

12x2-5-4 /xm. Paraphyses scanty, of two types: p.p. evenly distributed, branched, hyaUne, thin, 

0-7-1 pm wide, with apices sometimes widening to 2 ^tm but not becoming pigmented although 

embedded in pigmented gel-matrix; p.p. scattered or in small fascicles, mostly simple, stout, 2-3 

pm wide, coated ± throughout by dark pigment. Hypothecium 70-100(-160) pm tall, mottled, 

dark purplish brown, K-l- dark dull green, HNO3-I- purple-red; hyphae coated with densepigment 

and c. 1-5-3 pm wide, interwoven but becoming vertically orientated towards the 

hymenium and sometimes continuing into it as stout, pigmented paraphyses; ascogenous 

hyphae similarly pigmented but with short, swollen cells, to 4 pm wide. Excipulum indistinct- 

.


sometimes evident in young apothecia as a dark (concolorous with hypothecium) narrow zone 

forming an edge to the reflexed part of the hymenium; hyphae radiating, branched and 

anastomosing, c. 0-8-1 -5 /xm wide. 

Pycnidia numerous, black, sessile or more usually stalked, 60-300 jxm tall (including stalk) 

and 40-80 jxm diam; stalks (pycnidiophores) simple or sometimes branched bearing up to 4 

pycnidia; stalk and pycnidial wall tissues dark purplish brown, K+ dark green, HNO3+ 

purple-red. Conidiogenous cells A-A-1-6 /xm tall, with a cylindrical neck 1 •9-3-7 fxm tall and 

0-8-1-4 fxm wide, and with a swollen base 2-4-3(-3-7) /xm wide, the wall of which is often 

pigmented; percurrent proliferations not seen. Conidia (mesoconidia) ellipsoid or short cylin- 

drical, sometimes faintly biguttulate, 3-4-4-3x1-2-1-6 /xm. 

Chemistry: Thallus K- , C- , PD- ; apothecia sections C- ; no substances detected by t.l.c. 

Observations: Micarea nigella belongs to the group of lignicolous species (including M. 

contexta, M. eximia, M. melaeniza, M. misella, M. olivacea, and M. rhabdogena) with an 

endoxylic or indistinct thallus and small, ± globose to tuberculate, black apothecia. It is 

characterized by the purple-brown, K-h green pigment in the hymenium, hypothecium and 

pycnidial tissues, simple spores and stalked pycnidia. It is most likely to be confused in the field 

with other species with stalked, black pycnidia, namely M. melaeniza and M. misella. M. 

melaeniza differs in having a bright green hymenium, a red-brown hypothecium that does not 

turn green in K, a different pycnidial pigmentation and shorter mesoconidia. M. misella has a 

K-l- violet hymenium, a ± hyaline hypothecium and an olivaceous, K+ violet pigment in the 

pycnidial wall tissues. Apart from their lack of stalked pycnidia, M. contexta, M. eximia, and M. 

Map 15 Micarea nigella # -I- 

Micarea olivacea ® 

J


olivacea can be distinguished from M. nigella by their 1-septate spores. M. rhabdogena has a 

brown (K+ dissolving) pigment in the upper hymenium, and smaller spores. M. nigella could be 

confused with diminutive lignicolous forms of M. melaena, but that species usually has a bright 

green hymenium, larger and 1-3-septate spores, and never has stalked pycnidia. 

Habitat and distribution: M. nigella occurs on rather soft lignum of stumps of conifers and 

birch, in deeply shaded situations. Primarily, it appears to be an inhabitant of conifer 

woodlands, and has been found both in native pine forest (Black Wood of Rannoch) and mature 

conifer plantations. It is known from only three localities: north Jylland in Denmark, the 

southern central highlands of Scotland and north-east England; and, although undoubtedly a 

rather rare species, it has surely been much overlooked, at least in north-west Europe. 

30. Micarea nitschkeana (Lahm ex Rabenh.) Harm. 

(Fig. 24B;Mapl6) 

in Bull. Soc. Sci. Nancy II, 33: 64 (1899). - Bilimbia nitschkeana Lahm ex Rabenh., Lich. Europ. Exs. 

583 (1861). - Micarea denigrata var. nitschkeana (Lahm ex Rabenh.) Hedl. in Bih. K. svenska 

VetenskAkad. Handl. Ill, 18 (3): 79, 90 (1892). - Bacidia nitschkeana (Lahm ex Rabenh.) Zahlbr. in 

Annln naturh. Mas. Wien 22: 342 (1905). Type: Germany, Nordrhein-Westfalen, between Miinster and 

Nobriskrug, on Pinus in wood, T. R. J. Nitschke, Rabenh. Lich. Eur. Exs. 583 (M - lectotype!; 

isolectotypes: BM!, BM ex K!, M!). 

Lecidea spododes Nyl. in Flora, Jena 52: 410 (1869). - Bacidia spododes (Nyl.) Zahlbr. , Cat. lich. univ. 4: 

151 (1926). Type: England, Hampshire, Lyndhurst, New Forest, old pales, /. M. Crombie (H-NYL 

18819- lectotype!; isolectotypes: BM!, H-NYL 18820!). 

Bacidia nitschkeana f. microcarpa Erichsen in Verh. hot. Ver. Prov. Brandenb. 71: 97 (1929). Type: West 

Germany, Schleswig-Holstein, Eckernforde, near Levensau, Felmerholz, on Picea twigs, 9 xi 1924, 

C. F. E. Erichsen (HBG-holotype!). 

? Bilimbia spododes f. fusca B. de Lesd., Rech. Lich. Dunkerque: 200 (1910). Type: France, Nord, 

Dunkerque, Malo-Terminus, on dead branch of Hippophae, B. de Lesdain (not seen). 

?Bilimbia spododes f. livida B. de Lesd., Rech. Lich. Dunkerque, Suppl. I: 120 (1914). Type: France, 

Nord, Dunkerque, Zuydcoote, on dead branch oi Hippophae, B. de Lesdain (not seen). 

? Bilimbia spododes var. nigra B. de Lesd., Rech. Lich. Dunkerque, Suppl. I: 120 (1914). Type: France, 

Nord, Dunkerque, Malo-Terminus, dunes, on dried rhizome oiAmmophila arenaria, B. de Lesdain (not 

seen). 

Thallus effuse, usually forming small patches but sometimes wide-spreading, sometimes 

partly endoxylic but usually developed on the surface of the substratum as crowded, often 

contiguous, convex to subglobose areolae. Areolae especially well developed around the 

apothecia, dull greenish white to green-grey, surface matt, c. 40-200 /xm diam; in section, 

without a well defined cortex and not surrounded by an amorphous covering layer, outermost 

hyphae sometimes surrounded by dilute olivaceous, K-(- violet pigment. Thallus sometimes 

blackish and scurfy due to disruption by invading dematiaceous fungi and non-lichenized algae. 

Phycobiont micareoid, cells 4-7 /am diam. 

Apothecia scattered or, more usually, numerous and crowded, frequently contiguous or 

confluent, adnate, plane to convex-hemispherical, sometimes becoming tuberculate, immarginate 

or occasionally (especially when young) with an indistinct margin that is flush with the level 

of the disc, grey-black to black, or rarely whitish to pale grey-brown (shade forms), sometimes 

paler at the margin, matt. c. 0- 1-0-3 mm diam, or to 0-4 mm diam when tuberculate. Hymenium 

30-40 /xm tall, upper part dilute olivaceous to olivaceous, K+ violet, HNO34- red, C+ violet, 

also C-l- orange-red throughout due to gyrophoric acid; olivaceous pigment mostly confined to 

the gel-matrix, only rarely (in old, dark apothecia) closely bound to the walls of the paraphyses. 

Asci clavate 25-40x9-5-11 jxm. Spores fusiform, mostly curved, (l-)3(-4)-septate, 10-17 

(-19)x2-5-3(-3-5) pun. Paraphyses numerous, branched and anastomosing l-l-5(-l-7) jxm; 

apices not or only slightly incrassate, and mostly without closely adhering pigment. Hypothecium 

30-50 /am tall, hyaline; hyphae interwoven, c. 1-1-5 /am wide; ascogenous hyphae with 

swollen cells c. 2-5 /am wide. Excipulum distinct in sections of young, ± plane apothecia, but


reflexed and obscured in older, more convex apothecia; hyaline or dilute olivaceous (K+ violet) 

at outer edge; hyphae radiating, branched and anastomosing, c. 1-1-5 /jlui wide. 

Pycnidia indistinct but usually present, immersed in areolae; walls hyaline throughout, or 

olivaceous (K+ violet) around the ostiole; of three types: (a) c. 60-80 /xm diam, ostiole 

sometimes widely gaping; Conidia {macroconidia) curved or hamate, 1-3-septate, 12-26 xc. 1 

/Lim; (b) c. 50-100 (xm diam, ostiole often gaping and conidia extruded as a white blob; conidia 

(mesoconidia) short-cylindrical or obovate-oblong, apices rounded but often distinctly truncate 

at proximal end, sometimes biguttulate, (3-)3-5-5(5-7)xl-l-5 /xm; (c) c. 40-50 /xm diam, 

ostiole never gaping; conidia (microconidia) narrowly fusiform or bacilliform, (4-7-)5-5- 

7-5x0-7-0-8/>tm. 

Chemistry: Thallus K-, PD-; sections of thallus and apothecia C+ orange-red (also, parts 

with olivaceous pigment, C+ violet); t.l.c: gyrophoric acid. If thallus is heavily parasitized and 

scurfy the C+ orange-red reaction may be difficult to obtain and gyrophoric acid may not be 

detectable by t.l.c. 

Observations: M. nitschkeana is closely related, and similar in most respects, to M. denigrata 

{q.v.) with mainly 1-septate spores, and M. globulosella (q.v.) with ± acicular or rod-shaped 

0-3 (-6) -septate spores. Like M. denigrata, M. nitschkeana has three pycnidial anamorphs; 

usually only one or two of these states are found on any given thallus, but all three states (plus 

apothecia) have been noted on a single specimen from Scotland {Coppins 3369). In the field M. 

nitschkeana can be confused with species such as M. cinerea, M. lignaria, and M. peliocarpa, but 

these species usually have larger apothecia and areolae, and are quite different when examined 

microscopically. 

Habitat and distribution: M. nitschkeana is widely distributed in the British Isles at low 

altitudes (below 300 m), and is most frequently found on twigs or small branches of various 

deciduous and coniferous trees, e.g. Acer, Aesculus, Alnus, Betula, Fraxinus, Prunus padus, 

Quercus, Salix, Sambucus, Larix, and Picea; or on the old or dead stems of smaller shrubby 

plants, especially Calluna and Ulex, but also Erica cinerea, Myrica, Rosa, Sarothamnus, 

Viburnum, and Juniperus. It has been found to be abundant on old stems and Utter of Calluna in 

some lowland heaths in eastern England (Coppins & Shimwell, 1971), Scotland and Denmark, 

where it is commonly associated with Lecanora conizaeoides and Scoliciosporum chlorococum; 

such communities are referable to the Bacidietum chlorococcae . Other associated species on 

twigs and small branches include Lecanora pulicaris, L. symmicta agg., L. icmalea, Mycoblastus 

sterilis, Fuscidea lightfootii (western districts), Micarea prasina, Graphis elegans, Stenocybe 

pullatula (on Alnus) and small thalli of Hypogymnia physodes, H. tubulosa, Parmelia subaurifera, 

P. sulcata, Platismatia glauca, Evernia prunastri, and Ramalina fairinacea. It has 

occasionally been met with on lignum of fallen conifers (Larix and Pinus) or fence posts, habitats 

more characteristic of M. denigrata. Associated species on lignum include Lecanora confusa, L. 

symmicta agg. , Lecidea granulosa agg., L. icmalea, Mycoblastus sterilis, Hypogymnia physodes, 

Parmelia saxatilis, and Platismatia glauca. Occurrences on rock are rare, but a few collections 

have been made on sandstone or flint stones in heathland, with Lecidea granulosa agg., L. 

icmalea, and Micarea melaena as associates. It has also been found together with Scoliciosporum 

umbrinum on a plastic carton in a heathland in the Isle of Wight (Brightman & Seaward, 1977). 

M. nitschkeana is widely distributed through much of northern Europe. In Scandinavia it has a 

southern distribution and I have not seen material of it from north of latitude 62°N. It usually 

occurs in lowland situations, but it attains higher altitudes in the Alps (920 m) and the Nizke 

Tatry of Czechoslovakia (1250 m). Additional phorophytes from extra-British material include 

Hippophae (France: dunes near Calais), Vaccinium uliginosum (Germany: Schleswig- 

Holstein), and Pinus mugo subsp. pumilio (heathlands of southern Germany). From outside 

Europe I have seen only one specimen: USA, northern California (near San Francisco), where it 

occurred on fence ppsts. 

Exsiccata: Arnold Lick. Exs. Ill (BM ex K, M), 503 A, B (H-NYL, M); 503 C (BM ex K, H-NYL, M), 

503 D (H-NYL, M). Arnold Lick. Mon. 48 (BM ex K, M); 270 (BM ex K). Britz. Lick. Exs. 829 (M).


Map 16 Micarea nitschkeana # 1950 onwards O Before 1950 

Claudel & Harm. Lich. Gall. 89 (L, O). Harm. Lich. Loth. 853 (M). Hepp Flecht. Eur. 20 (BM, E, L, M), 

llp.p. (E). Krypt. Exs. Vindob. 1232 (M). Lojka Lich. Univ. 137 (BM ex K, M). Magnusson Lich. Sel. 

Scand. 340 (M). Malbr. Lich. Norm. 287 (M). Malme Lich. Suec. 25 (M, S). Rabenh. Lich. Eur. 582, 583 

(BM, BM ex K, M). Rasanen Lich. Fenn. 642 (M). Zahlbr. Lich. Rar. 110 (BM). Zwackh Lich. Exs. 470 

['bei Miinster in Westfalen'] (H-NYL 18825), 470 bis ['Bei Delbrueck in Kreise Paderborn.'] (H-NYL 

18822, M), 534 (H-NYL, M), 587 (H-NYL, M). 

31. Micarea olivacea Coppins, sp. nov. 

(Figs 24C, 47A; Map 15) 

Thallus effusus, endoxylicus vel epixylicus aut epilithicus, tenuissimus, inaequalis, interdum leviter 

areolatus, albidus vel viridio-griseus. Algae cellulis 4-7 ^tm diam. Apothecia immarginata, primum 

convexa vel subglobosa mox tuberculata, atra, 0- 1-0-3 mm diam, aut ad 0-4 mm diam ubi tuberculata. 

Hymenium 30-35 />tm altum, dilute sordide olivaceum cum vittis verticalibus atro-olivaceis, K-l- clare 

olivaceo-virescens. Ascosporae oblongae vel ovoideo-oblongae, rectae, (O-)l-septatae, (7-)9-12-3x2-5- 

3-5 jxm. Paraphyses numerosae, dimorphae: p.p. hyalinae, laxae, ramosae et anastomosantes, graciles, c. 

1-1-2 /xm latae, apicibus interdum leviter incrassatis ad 2 fxm latis; p.p. pigmentiferae, fasciculatae, 

plerumque simplices, crassae, 2-3 ^tm latae. Hypothecium sordide atro-olivaceum vel olivaceo-fuscum, 

K-l- virescens. Excipulum paulum evolutum mox reflexum. Pycnidia numerosa sed inconspicua, ± 

immersa, 25-50 ^tm diam., parietibus sordide olivaceis vel olivaceo-fuscis, K-l- clare olivaceis. Conidia 

brevitercylindrica, 3-4-2x1-1-3 />tm. Thallus ei apothecia K- ,C-, PD-. 

Typus: Caledonia, Mull, Aros, Druimfin, in ligno duro, 15 v 1968, P. W. James (BM - holotypus).


Thallus effuse, endoxylic to epixylic, or epilithic, forming a thin, whitish grey or greenish grey, 

uneven, sometimes weakly areolate crust; often appearing scurfy due to invasion by foreign 

fungi and algae. Phycobiont micareoid, cells 4-7 /xm diam. 

Apothecia numerous, immarginate, convex to subglobose, often tuberculate, black, matt or 

slightly glossy, surface minutely roughened, 0- 1-0-25 mm diam, or to 0-4 mm diam when 

tuberculate. Hymenium 30-35 /xm tall, without a distinct upper part (epithecium), dilute sordid 

olivaceous with dark olivaceous vertical streaks, K+ green intensifying, HNO3+ red. Asci 

clavate, 35-33x9-5-11 /xm. Spores oblong or ovoid-oblong, straight, (O-)l-septate, (7-) 

9-12-3x2-5-3-5 />im. Paraphyses numerous, of two types: p.p. hyaline throughout, evenly 

distributed, branched and often anastomosing, rather thin, 1-1-2 /^m wide, apices sometimes 

widening to 2 /am and sometimes with colourless, oily, refractive contents; p.p. broad, 2-3 /xm 

wide, mostly simple, grouped in small fascicles and often embedded in dense pigment. 

Hypothecium 40-70 /xm tall, dark sordid olivaceous or olive-brown, K+ green intensifying, 

HNO3+ red; hyphae embedded in greenish gel-matrix but walls not deeply pigmented, c. 1-5-2 

/am wide, interwoven but becoming vertically orientated towards the hymenium and sometimes 

continuing in to it as fasciculate paraphyses; ascogenous hyphae with short swollen cells, to 5 /am 

wide. Excipulum indistinct and soon reflexed, sometimes evident in sections of young apothecia 

as a hyaline or dilute sordid olivaceous, non-amyloid zone c. 20 /am wide; composed of radiating, 

much branched and anastomosing hyphae 0-8-1-5 /am wide. 

Pycnidia numerous but inconspicuous, ± immersed, 25-50 /am diam; walls sordid olivaceous 

or olive-brown, K-l- green intensifying, HNO3-I- red. Conidiogenous cells elongate-ampulliform 

to subcyUndric, 4—5-5x1-5-3 /am. Conidia (mesoconidia) short cylindrical, sometimes faintly 

biguttulate, 3-4-2x1-1-3 /am. 

Chemistry: Thallus K— , C— , PD— ; apothecia sections C— ; no substances detected by t.l.c. 

Observations: Micarea olivacea differs from M. eximia in its less brightly coloured hymenium 

pigmentation that is not concentrated in the upper part, more numerous and broader paraphyses, 

shorter and slightly broader spores with rounded apices, and shorter mesoconidia. From 

M. nigella it can be distinguished by the complete absence of purple pigmentation in water 

mounts, more numerous and broader paraphyses, more elongate and 1-septate spores, and ± 

immersed (never stalked) pycnidia. M. olivacea is easily confused with epruinose forms of 

the common lignicole Lecidea turgidula Fr., but that species has a dark green or olive-brown 

excipulum of conglutinated hyphae that do not separate in K, a paler hypothecium, and a thick 

walled, large celled phycobiont with cells 12-16 /am diam. In addition, the conidia of L. turgidula 

are smaller, c. 3-3-5x1-5-1-8 /am. When on rock M. olivacea can be confused with M. 

tuberculata, but the latter has a more brightly coloured hymenium and hypothecium, somewhat 

smaller spores, narrower asci, and a non-micareoid phycobiont with cells c. 5-10 /am when 

globose or up to 15 X 7 /am when ellipsoid . M. tuberculata has similar pycnidia and conidia , but its 

conidiogenous cells are more slender and about twice as long as those of M. olivacea. 

Habitat and distribution: M. olivacea is an apparently rare, but probably overlooked, species, 

being known from just two localities, both in Scotland. At the type locality it occurred on the 

hard Ugnum of a stump by a conifer plantation; the collection contains no associate species. At 

the other locality it occurred with Rhizocarpon hochstetteri and Baeomyces rufus on shaded rock 

in a mature conifer plantation. Little more can be said of its ecology until it becomes better 

known, but I am inclined to believe that it is essentially a lignicolous species with a preference for 

hard lignum . If this is true then care should be taken not to overlook it in the field for forms of the 

common M. denigrata with a reduced thallus.

32. Micarea osloensis (Th. Fr.) Hedl. 

(Fig. 28B) 


in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 83, 97 (1892). - Lecidea osloensis Th. Fr., Lich. 

Scand. 2: 524 (1874). Type: Norway, Oslo, summit of Ryenbjerget, on decaying plant debris, N. G. Moe 

(UPS-holotype! [t.l.c.: no substances]). 

Thallus effuse, indistinct, apparently of whitish grey, ± convex areolae, c. 50-70 /am diam, in 

the single specimen seen the thallus is obscured by a bleached non-lichenized alga, giving it a 

pale grey 'filmy' appearance. Phycobiont ^xohdhXy micareoid, cells 4-5-6-5 /am diam. 

Apothecia numerous, ± evenly scattered, immarginate, convex to subglobose, black or 

brown-black, 0- 1-0-26 mm diam. Hymenium c. 30 /am tall; upper part (epithecium) red-brown, 

K— , HNO3— 

or red tinge intensifying; remaining (lower) part dilute reddish brown with darker, 

red-brown, vertical streaks. Asci clavate, 26-30x 11-13 /am. Spores ellipsoid or ovoid-ellipsoid, 

simple, 6-9-5x3-4 /am. Paraphyses rather numerous, simple below, but mostly branched 

towards their apices, hyaline and 1-5-1-8 /am wide, but occasionally surrounded throughout 

their length by red-brown pigment and then 1-7-2 /am wide; apices sometimes swollen to 3 /am 

wide, surrounded by an amorphous, red-brown, pigmented matrix that does not disperse in K. 

Hypothecium c. 70-85 /am tall, dark reddish brown, K— , HNO3— 

Excipulum not evident, even in sections of young apothecia. 

Pycnidia not seen. 


detected by t.l.c. 

, PD— 

or red tinge intensifying. 

; sections of thallus and apothecia C— ; no substances 

Observations: Micarea osloensis is characterized by its small, blackish apothecia with internal 

reddish brown pigment that does not disperse in K, simple, ellipsoid spores, and terricolous 

habitat. In the field it is most likely to be overlooked for Lecidea uliginosa, but that species has a 

well developed, pseudoparenchymatous excipulum, paraphyses with a brown apical cap, larger 

spores, and 'Trapelioid' asci which do not have a deeply amyloid tholus. 

Habitat and distribution: Knowledge of M. osloensis is still restricted to its type gathering from 

Norway. The appearance of this material suggests that it occurred in a woodland clearing on the 

site of an old bonfire (small amounts of ash are present), probably in a slight depression. The 

specimen is accompanied by the sterile thallus of Lecidea icmalea and a few plants of the moss 

Ceratodon purpureas. Dr H. Krog informs me that the hill Ryenberget is now well within the city 

limits of Oslo, and is now much affected by the results of urbanization, although not completely 

destroyed. It has an altitude of about 150-200 m, and was almost certainly covered in pine forest 

before it was engulfed by the city. 

33. Micarea peliocarpa (Anzi) Coppins & R. Sant. 

(Figs 3A-C, 25, 48, 53-54; Map 17) 

in Coppins & P. James in Lichenologist 11: 155 (1979). - Bilimbia peliocarpa Anzi in Atti Soc. ital. Sci. 

nat. 9: 250 (1866). - Bacidia peliocarpa (Anzi) Lettau in Hedwigia 52: 133 (1912). - Micarea violacea f. 

peliocarpa (Anzi) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 81, 92 (1892); nom. superfl. 

(Art. 63). Type: Italy, Piemonte, Novara, 'Sopra i muschi nei monti del Lago Maggiore (Locarno), 

scoperta dal padre capuccino Agostino Daldini' (? TO, not seen: request for loan not acknowledged); an 

unlocalized specimen labelled 'Bilimbia peliocarpa Anzi Neosymb. no. 44' [reference to original 

protologue] in Anzi's handwriting occurs in UPS(!) and is possibly an isotype. 

Biatora lignaria (3. [var.] conglomerata Hepp, Flecht. Eur., fasc. 5, no. 284 (1857). - Micarea violacea f. 

conglomerata (Hepp) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 81, 92 (1892). Type: 

Switzerland, 'bei Rifferschweil', on Finns bark, Hegetschweiler, Hepp Flecht. Eur. 284 (E - lectotype!; 

M-isolectotype!). 

Biatora lignaria var. saxigena Hepp, Flecht. Eur. fasc. 9, no. 510 (1860). Type: Germany, Hessen, 'auf 

Sandsteinfelsen bei Marburg', W. Uloth, Hepp Flecht. Eur. 510(E-lectotype!; M-isolectotype!). See 

note (i) below.


Bilimbia milUaria var. lignaria *[f.] livida Korber, Parerga Lich.: 171 (1860). Type: same as Biatora 

lignaria var. conglomerata Hepp. 

Lecidea sphaeroides var. leucococca Nyl. in Stizenb. in Nova Acta Leop. - Carol. 34 (2): 18, 1. 1 , f. D, 47-51 

(1867). Type: Finland, Tavastia australis, Evois ['Evo'], on lignum, 1865, J. P. Norrlin (H-NYL 18377lectotype!; 

H-isolectotype!). 

Bilimbia violacea Arnold in Flora, Jena 53: 473 (1870). - Lecidea violacea P. Crouan & H. Crouan ex Nyl. 

in Flora, Jena 45: 464 (1862), non Massal. (1852). - Micarea violacea (Arnold) Hedl. in Bih. K. svenska 

VetenskAkad. Handl. Ill, 18 (3): 80, 91 (1892). Type: France, Finistere, Brest, on schistose rock, 

Crouan (H-NYL 18716- isotype!). [Note: listed as 'Bacidia violacea (Crouan ex Nyl.) Arnold' by James 

(1965a); this should not be confused with Bacidia violacea (Arnold) Arnold, Flora, Jena 67: 581 (1884), 

which is a Bacidia s. str.] 

Bilimbia trisepta Hellb., Nerikes Lafflora: 11 (1871). - Biatora trisepta Naeg. ex Mull. Arg. in Mem. Soc. 

Phys. Hist. nat. Geneve 16: 403 (1862); as 'Naeg. mss. ex Dr Hepp', nom. inval. (Arts 32, 34). - Lecidea 

sabuletorum f. trisepta Stizenb. in Nova Acta Acad. Leop. -Carol. 34 (2): 47, t. 3, f. A, 35-62 (1867); nom. 

superfl. (Art. 63). - Bilimbia hypnophila b. [var.] trisepta Bausch in Ver. naturw. Ver. Karlsruhe 4: 127 

(1869); nom. superfl. (Art. 63). - Bilimbia sabuletorum d. [var.] trisepta Rabenh., Krypt.-Fl. Sachsen 2: 

187 (1870); nom. superfl. (Art. 63). - Bacidia trisepta (Hellb.) Zahlbr. in Engler & Prantl, Nat. 

Pflanzenfam. 1 (1*): 135 (1905). -Micarea trisepta (Hellb.) Wetmore in Pub. Mus. Mich. St. Univ. Biol. 

3: 284 (1968). Type: Germany, Hessen, 'auf Sandsteinfelsen bei Marburg', W. Uloth, Hepp Flecht. Eur. 

510 (E-neotype!; M-isoneotype!). See note (ii) below. 

Lecidea hemipolioides Nyl. in Flora, Jena 56: 294 (1873). - Bilimbia hemipolioides (Nyl.) A. L. Sm., 

Monogr. Br. Lich. 2: 141 (1911). - Bacidia hemipolioides (Ny\.) Zahlbr. , Cat. lich. univ. 4: 114(1926).- 

Micarea violacea f. hemipolioides (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 80, 91 

(1892). Type: Channel Islands, Jersey, Rozel meadow, on rocks, 1873, C. Larbalestier (H-NYL 18713 - 

lectotype!; isolectotypes: H-NYL p.m. 4613!, BM!). 

Bilimbia naegelii f. lapiseda Th. Fr., Lich. Scand. 2: 379 (1874). Type: Sweden, Stockholm, Lyran, on 

rock, 1870, S. Almquist (IJ?S - lectotype!). 

Bilimbia milliaria f. livescens Th. Fr., Lich. Scand. 2: 383 (1874). Type: Sweden, Narke, Gotlunda, 'pa 

stenmuren mellan Stabacken och Hogby', on mosses, 1863, Blomberg (UPS - lectotype!). 

Lecidea albidolivens Nyl. in Flora, Jena 57: 10 (1874). - Bilimbia albidolivens (Nyl.) Blomb. & Forss., 

Enum. PI. Scand.: 82 (1880). Type: Finland, Tavastia australis, Padasjoki, Nyystola, on lignum, 1872, 

E. A. Lang fVainioj (H-NYL 18775 - lectotype!; H - isolectotype!). 

Lecidea fraterculans Nyl. in Flora, Jena 58: 11 (1875). Type: Finland, Tavastia australis, Padasjoki, 

Nysstola, on rock, 1872, E. A. Lang [VainioJ (H-NYL 18704a - lectotype!; H-NYL 18704 - isolecto- 

type!). 

Lecidea triseptatula Nyl. in Flora, Jena, 58: 361 (1875). Type: Finland, Tavastia australis, HoUola, on 

lignum, 1874, E. A. Lang fVainioj (H-NYL 18709 -lectotype!; isolectotypes: H-NYL 18710!, H!). 

Bilimbia albicans Arnold in Flora, Jena 65: 140 (1882). Type: Germany, Bayern, Oberbayern, near Bad 

Tolz, W side of the Blombergs, on sandstone in wood, 5 ix 1880, Arnold, Lich. exs. 837 (BM - 

lectotype!). 

Micarea violacea f. cupreola Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 80, 91 (1892). Type: 

Sweden, Halsingland, Jarvso, on rock, viii 1891, Hedlund(S-\\o\oiypQ\). 

Lecidea triseptatuloides Harm, in Bull. Seanc. Soc. Sci. Nancy II, 33: 64 (1899). - Bacidia triseptatuloides 

(Harm.) Zahlbr., Cat. lich. univ. 4: 161 (1926). Type: France, Moselle, Bitche, on Pinus bark, xii 1893, 

Abbe Kieffer (ANGUC - holotype!). 

Micarea violacea f. exigua Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 81, 91 (1892); nom. 

inval. (Art. 26). 

Notes: (i) 'saxigena'. This epithet has been applied at intra-specific levels to saxicolous forms of M. 

lignaria and M. peliocarpa. Its first valid publication was by Hepp (loc. cit.) as "Biatora lignaria (Ach.) var. 

saxigena (Leight) Hepp' with 'Lecidea milliaria (Fries) v. saxigena, Leight Lich. Brit. Exs. 210' given as a 

synonym. All examples seen by me of Leighton Lich. Brit. 210 (issued in 1856) are M. lignaria; whereas all 

examples of Hepp Flecht. Eur. 510 are M. peliocarpa. Hepp's illustration and description of spores are 

more applicable to M. peliocarpa than M. lignaria. The original valid introduction of the epithet "saxigena' 

should, therefore, be typified on an example of Hepp Fl. Eur. 510 and attributed to Hepp alone. 

Theoretically "saxigena Leighton' could have been validated for saxicolous forms of M. lignaria by 

providing a description, excluding other names or specimens that would render it superfluous, and 

avoiding homonymy with previous combinations of "saxigena Hepp' ; but I have not found such a case in the 

Hterature. 

(ii) "trisepta". This epithet first appeared as "Biatora trisepta Naeg. mss. ex Dr Hepp' in the protologue of


Patellaria salevensis Miill. Arg. (= Bacidia salevensis (Mull. Arg.) Zahlbr.). Miiller presumably regarded 

Naegeli's manuscript name as a synonym of his new species. B. salevensis (type in BM!) is a limestone 

species and is not a Micarea. As indicated by James (1965: 103) the first legitimate publication of the epithet 

'trisepta' for a Bacidia-\\kt species is that by Hellbom {loc. cit). By including the epithet in Bilimbia as '5. 

trisepta (Naeg.)', Hellbom was clearly using it in its traditional sense and not introducing it as an entirely 

new name. There is no indication that any of the authors using the epithet 'trisepta Naeg.' had seen 

Naegeli's original material, and no such material has been found amongst Hepp's collections at BM. 

Because the original material is probably no longer in existence, neotypification is required. The epithet 

'trisepta'' has been a long source of confusion, being used by various authors for several species, especially 

Micarea lignaria, M. melaena, M. nitschkeana, M. peliocarpa, and M. ternaria. However, it is clear from 

the descriptions and cited specimens in Stizenberger (loc. cit.), Bausch (loc. cit.), Hedlund (1892), and 

Vainio (1922) that the epithet 'trisepta' has most commonly been applied to the species now called Micarea 

peliocarpa. Because 'trisepta' first appeared in the protologue of a saxicolous species the neotype chosen is 

an example of Hepp Flecht. Eur. 510 which was cited under, or in connection with, 'trisepta' in the 

treatments of Stizenberger, Bauch, Hedlund, and Vainio. 

Thallus effuse, sometimes partly endocuticular, endophloeodal or endoxylic, more usually 

developed on the surface of the substratum as rounded, shallow-convex, hemispherical, or ± 

globose areolae. Areolae scattered or more usually ± continguous, greenish white, or greywhite 

to blue-grey, occasionally becoming dark grey, often dark coloured on upper surface but 

pale and greenish white below, matt or slightly glossy, c. 40-200 ixm diam. Areolae in section, 

ecorticate but with an amorphous hyaline covering layer c. 2-5 /x,m thick; outermost hyphae 

often with grey-green to blue-green walls, K-, HNO3+ red. Phycobiont micareoid, cells 4-7 

/xm diam. 

Apothecia scattered or more usually crowded and often contiguous, adnate, plane to convex, 

sometimes becoming tuberculate; sometimes with an indistinct margin that is flush with the level 

of the disc; rarely (as in shaded corticolous forms) whitish or ivory-white, usually pale lead-grey 

to grey-black, sometimes grey-brown, often whitish or paler at the margin; (0-12-)0- 14-0-4 

(-0-6) mm diam, or to 1 mm when tuberculate. Disc finely roughened, matt, or slightly glossy 

(when black); margin (when evident) smooth and often more glossy than the disc. Hymenium 

40-55 iJLm tall, usually olive-green or aeruginose (K— , HNO3-I- red) in upper part and ± hyaline 

below; in dark coloured apothecia the green pigment often occurs as vertical streaks through the 

hymenium. Asci clavate c. 40-55x12-17 /xm. Spores fusiform, clavate-fusiform, or oblongfusiform, 

often slightly curved, (l-)3(-5)-septate, (ll-)15-23(-24)x3-5(-6) /am. Paraphyses 

numerous, branched, often anastomosing, 1-1-5 /xm wide; apices often more richly branched 

and entangled, often slightly incrassate to c. 1-8 /xm, or to 2-5 /i,m due to thickening by green 

pigment. Hypotheciumc. 40-70 />im tall, hyaUne or dilute straw; hyphae interwoven, c. 1-1 -7 /xm 

wide; ascogenous hyphae with swollen cells c. 2-4 /xm wide. Excipulum well developed, hyaline, 

or dilute straw in part, of richly branched and anastomosing, radiating hyphae c. 1-1-5 îm. 

Pycnidia frequently present (especially on bark or lignum), of two types: (a) immersed in 

areolae, white or greenish around the ostiole, 140-200 /xm diam, ostiole often widely gaping; 

conidia (macroconidia) markedly curved and often sigmoid, often faintly 1-5-septate, 21-40(- 

50)x 1-1-5 fim; (b) ± sessile, white, 50-80 /xm diam, ostioles not, or only slightly, gaping; 

conidia (microconidia) narrowly fusiform-cylindrical, (5-)6-7(-7-7)x0-4-0-7 /xm. 

Chemistry: Thallus and whitish apothecia K-, C-lorange-red; 

t.l.c: gyrophoric acid. 

red, PD— ; apothecia in section C-l- 

Observations: M. peliocarpa occurs in a wide range of habitats and is concurrently variable, 

especially regarding the colour of its apothecia and thallus. This variation is considered to be 

phenotypic, being due to the amount of green pigment produced in the hymenium and near the 

surface of the thallus in response to environmental factors, particularly light. When on the bark 

of old trees in sheltered woodland the apothecia are often very pale to blue-grey, more scattered 

than usual, and accompanied by numerous pycnidia; such forms are frequent in the ancient 

woodlands of the New Forest, Hampshire. There is some variation as to the production of 

areolae, these being often poorly developed or even absent when on cortex or lignum, and


usually abundant and well developed in more exposed situations on mossy rocks and peaty 

debris. 

M. peliocarpa is most closely related to M. alabastrites and M. cinerea (see under the former 

for further discussion). Apart from these two species, M. peliocarpa is most often confused with 

M. lignaria {q.v.) especially when occurring in exposed habitats where its apothecia may be 

black and markedly convex. Similarly, confusion could occur with the rare M. ternaria {q.v.), 

which like M. peliocarpa has rather flattened apothecia (in section) with a well developed 

excipulum. When fertile, M. leprosula has apothecia very hke those of M. peliocarpa, but its 

thallus is composed of fragile areolae that readily dissolve into soredia and contains argopsin 

(PD+ red) as well as gyrophoric acid. In certain habitats, such as the stems of old shrubs (e.g. 

Calluna, Erica, and Ulex), M. peliocarpa could be mistaken for M. nitschkeana although the 

latter has an olivaceous, K+ violet pigment in the hymenium, pycnidial walls and thallus, and 

smaller spores. Specimens collected from Xanthorion communities and attributed to M. 

peliocarpa (or one of its synonyms) are mostly referable to Bacidia naegelii, which has simple 

paraphyses with markedly swollen apices, an excipulum of coherent hyphae (in K), and a very 

different thallus structure. 

Habitat and distribution: M. peliocarpa is rather catholic in its choice of habitat. As an 

epiphyte on bark (or on bryophytes thereon) it is mostly commonly found on mature Quercus, 

but has also been collected from Alnus, Betula, Fagus, Fraxinus, Ilex, Salix, Larix, Pinus, 

Juniperus, Calluna, Erica, and Ulex. It has a preference for trunks and main stems rather than 

small twigs and branches. Amongst the British collections associated lichens on bark include 

Arthonia spadicea, Biatorina atropurpurea, 'Botrydina vulgaris', Cladonia spp., Hypogymnia 

physodes, Lecanora pallida, L. symmicta agg., Lecidea icmalea, Leparia incana agg., Micarea 

alabastrites, Micarea cinerea, Normandina pulchella, Ochrolechia androgyna, Farmelia saxatilis, 

Parmeliella jamesii, Pertusaria hymenea, Phyllopsora rosei, Platismatia glauca, Stenocybe 

septata and Trapelia sp. In the north and west of Britain it is frequently encountered on the 

lignum of fallen trunks and large branches (especially conifers), accompanied by such species as 

Lecidea granulosa agg. , L. turgidula, Micarea denigrata, M. lignaria (including var. endoleuca), 

Mycoblastus sterilis, Ptychographa xylographoides, Xylographa abietina, and X. vitiligo. In the 

same areas it is often common on peaty soil, and on moribund bryophytes or peaty debris on old 

walls, boulders and rock faces, preferring rather more sheltered conditions than M. lignaria, 

with which it is easily confused in the field. Furthermore, it is not found at such high altitudes as 

attained by that species, and all British collections seem to have been made at below 500 m. 

Associated species in these habitats include Cladonia crispata, C. coccifera, C. squamosa, C. 

uncialis, Coelocaulon aculeatum s. lat., Hypogymnia physodes, Lecidea icmalea, Micarea 

leprosula, Ochrolechia androgyna, Farmelia saxatilis, dind Platismatia glauca. It is less frequent- 

ly found growing directly on rock, and records from Britain indicate that it is able to do so only in 

rather dry situations. Such occurrences are mostly in the rather low rainfall districts of, for 

example, Durham, east Yorkshire and Sussex; accompanying species noted include Baeomyces 

rufus, Cystocoleus ebeneus, Lecidea granulosa agg. , L. icmalea, Lecanora polytropa, Lepraria 

incana agg., Farmeliopsis ambigua, Trapelia coarctata, and T. involuta. 

M. peliocarpa is widely distributed in Britain but is most common in the west and in upland 

districts (but at low altitudes). This pattern is reflected in Europe as a whole, and it is apparently 

common in countries along the Atlantic seaboard and on the Atlantic islands including Iceland, 

the Azores, and Canary Islands. In central and eastern Europe it is found mainly in mountainous 

districts. Northwards it extends to just beyond the Arctic Circle, but I do not know of its 

occurrence in the high arctic. From outside Europe I have seen material from eastern Canada 

and north-eastern USA, and from New Zealand. 

Exsiccata: Anzi Lick. Sondr. 170A (UPS). Arnold Lich. Exs. 167A, B (BM ex K, M), 837, 1051 (BM ex 

K). Arnold Lich. Mon. 1 18 (BM ex K), 269, 357, 482 (BM ex K, MANCH). Hepp Flecht. Eur. 284 (E, M), 

285 (E, L, M), 510 (E, M). Hepp Zur. 206 (BERN). Korber Lich. Sel.Germ. 133A, B (M). Krypt. Exs. 

Vindob. 165 (BM). Larb. Lich. Herb. 347 (BM). Leighton Lich. Brit. 23^ p.p. (BM, DBN, FRS). Lojka 

Lich. Hung. 134 (BM ex K). Malme Lich. Suec. 169 (S). Rasanen Lichenoth. Fenn. 343 (BM). Vezda Lich. 

Sel. 1342, 1380 (BM). Zwackh. Lich. Exs. 276, 897 (UPS).

34. Micarea prasina Fr. 


-^^r 

Map 17 Micarea peliocarpa # 1950 onwards O Before 1950 

(Figs ID, 26-27, 49, 53-54; Maps 18-20) 

Syst. orb. : 256-7 (Dec. 1825). - Biatora prasina Fr. , Stirp. agrifemsion.: 36 (June 1825), nom. illeg. (Art. 

63). - Biatora prasina (Fr.) Trevisan, Linnaea 28: 288 (1856), nee Tuck. & Mont., in Mont. (1857). - 

Catillaria prasina (Fr.) Th. Fr., Lich. Scand. 2: 572 (1874). See note (i) below and note on Sporacestra 

under 'Excluded taxa'. 

Biatora micrococca Korber, Parerg. lich.: 155 (1860). - Catillaria micrococca (Korber) Th. Fr., Lich. 

Scand. 2: 571 (1874). - Micarea prasina f. micrococca (Korber) Hedl. in Bih. K. svenska VetenskAkad. 

Handl. Ill, 18 (3): 77, 87 (1892). - Micarea micrococca (Korber) H. Gams, Kleine Kryptfl. 3: 67 (1967); 

comb, inval. (Art. 33.2). Type: Germany, Baden-Wiirttemberg, 'in regno Wurtembergico', on Pinus 

bark, 1862, K. A. Kemmler, Korber Lich. Sel. germ. 250 (L 910, 139-1361-neotype! [t.l.c: 'prasina- 

unknown A']). See note (ii) below. 

Lecidea subviridescens Nyl. in Flora, Jena 51: 474 (1868) . - Micarea subviridescens (Nyl.) Hedl. in Bih. K. 

svenska VetenskAkad. Handl. Ill, 18 (3): 77, 87 (1892). - Bacidia subviridescens (Nyl.) Zahlbr., Cat. 

lich. univ. 4: 154 (1926). Type: Channel Islands, Jersey, Boulay Bay, on coastal turf and soil, 1868, C. 

Larbalestier (H-NYL 19056 [as 'subvirescens'] - lectotype! ; BM - isolectotype! [t.l.c. : 'prasina unknown 

C']; BM-topotypes!). See note (iii) below. 

Lecidea prasiniza Nyl. in Flora, Jena 57: 312 (1874). - Micarea prasina f. byssacea subf . prasiniza (Nyl.) Th. 

Fr. in Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 77 (1892). Type: Finland, Tavastia 

australis, Padasjoki, Nyystola, on Alnus, 1872, E. A. Lang fVainio] (H-NYL 21604 - lectotype!, H - 

isolectotype!). [material insufficient for t.l.c.]. 

Lecidea sordidescens Nyl. in Flora, Jena 57: 312 (1874). - Lecidea erysiboides f. sordidescens Nyl. in 

Norrlin in Not. Sdllsk. Fauna Fl. fenn. Forh. 11: 188 (1870); nom nudum (Art. 32). - Catillaria prasina


var. byssacea f . sordidescens (Nyl.) Blomb. & Forss. , Enum. PI. Scand. : 91 (1880). - Micarea prasina f 

byssacea subf. sordidescens (Nyl.) Th. Fr. in Hedl. in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 

77 (1892). - Lecidea byssacea var. sordidescens (Nyl.) Vainio in Termeszetr. Fuz. 22: 320 (1899). - 

Catillaria prasina var. sordidescens (Nyl.) Lettau in Hedwigia 52: 136 (1912). - Micarea prasina var. 

sordidescens (Nyl.) Brodo in Bull. N. Y. St. Mus. Sci. Serv. 410: 152 (1968). Type: Switzerland, Zurich, 

'An faulen Fichten-Strunken Z. H.', Hepp, Flecht. Eur. 11^ (E - lectotype! [t.l.c.: 'prasina-unknown 

B']; isolectotypes: BM (boxed set)! [t.l.c: 'prasina-unknown B'], H-NYL 21632 [fragment]!, M!). See 

note (iv) below. 

Lecidea prasiniza var. prasinoleuca Nyl. in Flora, Jena 64: 7 (1881). Type: Germany, Baden-Wiirttemberg, 

Heidelberg, Konigstuhle, on Picea abies, ix 1880, Zwackh, Lich. Exs. 593a (H-NYL 21601 - lectotype! 

[t.l.c: 'prasina-unknown A']). See note (v) below. 

Catillaria micrococca f. glebosula Erichsen in Annls mycol. 36: 139 (1938). Type: Germany, Schleswig- 

Holstein, Eiderstedt, Ording, at base of Betula, 4 viii 1913, C. F. E. Erichsen (HBG - holotype! [t.l.c: 

'prasina-unknown B']. 

Catillaria micrococca var. discrepans Erichsen in Annls mycol. 36: 139 (1938). Type: Denmark, Jylland, 

Abenra (Apenrade), on smooth bark oiAlnus in a valley near Elisenlund, 3 ix 1913, C. F. E. Erichsen 

(HBG - holotype!). [material insufficient for t.l.c]. 

Catillaria prasina var. occulta Erichsen in Annls mycol. 36: 140 (1938). Type: Germany, Schleswig- 

Holstein, Ratzeburg, in the wood 'Bak', on ± smooth bark oiAlnus, 21 iii 1918, C. F. E. Erichsen (HBG 

- holotype!). [material insufficient for t.l.c]. 

Lecidea declivitatum Erichsen in Mitt. Inst. allg. Bot. Hamb. 10: 408 (1939). Type: Germany, Schleswig- 

Holstein, 'Krs Eutin: an der Boschung eines Waldwegs an der Nordwest-Seite des Ugleisees', 5 vi 1914, 

C. F. E. Erichsen (HBG -holotype! [t.l.c: 'prasina-unknown C']). 

Micarea polytrichi Poelt & Dobb. in Bot. Jb. 96: 343 (1975). Type: Austria, Steiermark, Grazer Bergland, 

a little north of Maria Trost near the Wenisbucher Strasse, c. 460 m, on Polytrichumformosum on shady 

side of a narrow gorge, 30 iii 1974, /. Poelt (GZU - holotype!) [material insufficient for t.l.c]. 

Catillaria prasina a laeta Th. Fr., Lich. Scand. 2: 573 (1874); nom. inval. (Art. 26). 

?Biatora byssacea Zwackh in Flora, Jena 45: 510 (1862), non Hampe in Linnaea 25: 709 (1852). Type: 

Germany, Heidelberg, 'Konigsstuhle', on bark of old Quercus, ? Zwackh (not traced). See note (vi) 

below. 

?Catillaria melanobola f. biseptata B. de Lesd., Rech. Lich. Dunkerque: 198 (1910). Type: France, Nord, 

Dunkerque, St Pol, on a piece of leather in dunes, B. de Lesdain (not seen). 

?Catillaria melanobola f. nigra B. de Lesd., Rech. Lich. Dunkerque: 199 (1910). Type: France, Nord, 

Dunkerque, lignum of hollow Salix, B. de Lesdain (not seen). 

Notes: (i) Biatora prasina Fr. (June 1825) is superfluous because the protologue includes the sentence: 

'Crusta est Byss. botryoides Linn!'. Byssus botryoides L. is a valid name {Sp. pi. 2: 1169 (1753)) but the 

material in the Linnaean herbarium (LINN) is a specimen (1278.16) labelled by Ehrhart; it is an alga and 

bears a determination label: 'Protococcus viridis Ag. Determined by Francis Drouet iv. 1950'. In the 

protologue of Micarea and M. prasina. Fries (Dec. 1825) makes a less dogmatic statement, merely 

indicating that when sterile M. prasina is indistinguishable from B. botryoides; M. prasina is, thereby, not 

rendered superfluous. 

(ii) I have seen two specimens named by Korber and collected from 'Wiirtemberg' by Kemmler: an 

undated specimen in WRSL and possibly part of the original gathering, and a later specimen issued in 

Korber's exsiccata. The first specimen would have to be completely destroyed for t.l.c. analysis, whereas 

the exsiccate specimen is larger and was found to contain 'prasina-unknown A', and it is this specimen that I 

have chosen for typification. If future studies require this chemical race to be recognized at specific rank, 

then 'micrococca' is the earliest available epithet. 

(iii) L. subviridescens is the earliest name for the chemical race with 'prasina-unknown C. 

(iv) Although Nylander (1874) did not provide a description, he validated L. sordidescens by referring to 

it Hepp Flecht. Eur. 278; the printed label for this exsiccatum includes a short description and a drawing of 

spores. 

(v) If the chemical race containing 'prasina-unknown A' were to be awarded varietal rank, then this 

name provides the earliest available epithet, provided that 'byssacea' is not available; see also note (vi). 

(vi) I have not seen a holotype specimen or any material suitable for the lectotypification of Biatora 

byssacea Zwackh. A neotype has not been selected because this name is a later homonym of B. byssacea 

Hampe. Hampe's name was given to a specimen collected by Charles Stuart in Tasmania, and the 

protologue ends with 'an status degenerans'. Biatora byssacea Hampe is included by Zahlbruckner (Cat. 

lich. univ. 3: 897) in a list of excluded and dubious names and is not accorded a catalogue number; the type 

.


material has not been located in BM but it could be elsewhere (? G). Under the taxonomic arrangement 

given here Zwackh's epithet is of no consequence. If the chemical races of M. prasina were to be awarded 

varietal status, then under the provisions of Article 72 the name ' Catillaria prasina p. [var.] byssacea Th. Fr. 

(1874)' could become important and require typification. If this name was typified with a specimen 

containing 'prasina unknown A' then it would be the earliest available name at that rank. 

Thallus effuse and often wide-spreading, very variable in colour, pale fawn (dry shaded 

situations), light green, pale to dark grey-green (often with glaucous hue), or olivaceous to 

olive-black, often appearing ± gelatinous when moist, composed of small ± globose granules 

(goniocysts). Goniocysts c. 12^0(-60) jxm diam, thinly scattered, or vertically proliferating to 

give the thallus a soft isidiose appearance, or densely aggregated to form a thick granular crust 

up to 200 />tm deep. Goniocysts ecorticate, often with shortly protruding hyphal filaments; 

outermost hyphae of superficial goniocysts sometimes surrounded by greenish, K+ violet 

pigment. Phycobiont micareoid, cells 4-7 ^tm diam. 

Apothecia usually numerous but sometimes few or absent, immarginate, sometimes adnate 

and shallow-convex, but more usually becoming markedly convex, subglobose or tuberculate; 

colour variable, whitish or pallid {'micrococca' shade-forms), pale to dark grey, brownish grey, 

or grey-black, matt; 0- 1-0-4 mm diam, or to 0-6 mm when tuberculate. Hymenium 28-45(-52) 

H^va tall, hyaline or dilute dull straw, but more usually dull olivaceous, especially in the upper 

part (but coloured portion not sharply delimited) or in vertical streaks; pigment K+ violet, 

HNO3+ red, confined to gel-matrix. Asci clavate or cylindrical-clavate, 26-40(-50)x8-12 /u,m. 

Spores ovoid-ellipsoid, ovoid, ovoid-oblong or oblong, 0-l(-3)-septate; when 1-septate the 

lower cell is slightly longer and narrower than the upper, and there is often a slight constriction at 

the septum; variable in size, in range 8-14(-17)x2-3-4(-5) /xm. Paraphyses rather numerous, 

branched and anastomozing, 0-5-1 /xm wide; apical parts often wider, to 1-5 /xm, but not 

thickened by pigment; epithecial pigment confined to surrounding gel matrix. Hypothecium c. 

40-170 /xm tall, hyaline to dilute dull yellowish; hyphae interwoven, some becoming vertically 

orientated towards the hymenium, c. 0-7-1-3 jxm wide; ascogenous hyphae with swollen cells, c. 

2-4 /xm wide. Excipulum poorly developed, sometimes evident as a narrow, hyaline or dilute 

olivaceous-straw (K-l- violet), non-amyloid, reflexed zone; hyphae radiating, branched and 

anastomosing, c. 0-7-1 /am wide. 

Pycnidia often present but usually inconspicuous, white, or upper part around the ostiole grey 

(due to olivaceous, K+ violet pigment), immersed to sessile (never distinctly stalked), of two 

types: (a) c. 50-120 /xm diam, emergent to sessile, ostiole sometimes gaping; conidia (mesoconi- 

dia) ± cylindrical or narrowly obpyriform, often biguttulate and slightly constricted near the 

middle, (3-5-)4-6x 1-2-1-7 /xm; (b) c. 30-60(-100) /xm, usually immersed in surrounding 

goniocysts, ostiole rarely gaping; conidia (microconidia) cylindrical or narrowly fusiform, 

(5-)5-5-8x0-7-l/xm. 

Chemistry: All parts K— , C— 

(but olivaceous pigment C-l- violet, never red), PD— ; t.l.c: 

specimens have one of three unknowns ('prasina unknown A', 'B', or 'C; see p. 87); trace 

amounts of gyrophoric acid sometimes detected, possibly due to contamination by intimately 

associated species such as Lecidea icmalea and L. granulosa agg. 

Observations: Micarea prasina is the commonest and most variable member of the genus. 

However, it is one of the few European species whose thallus is composed of minute discrete 

granules (goniocysts). Two other species with a similar thallus are: M. hedlundii (q.v.) with a 

K-l- red oily substance in the goniocysts and conspicuous tomentose, brown, stalked pycnidia; 

and M. melanobola which has a sharply delimited epithecium (pigment closely adhering to 

apices of the paraphyses) and shorter microconidia . Both of these species appear to be very rare 

A comparison of the diagnostic features of M. melanobola, M. prasina and M. misella are given 

in Table 6. 

Diminutive forms of M. prasina with small pallid apothecia and a scanty thallus have 

sometimes been regarded as a distinct species {'Catillaria micrococca') . 

Such forms have the 

same basic thallus structure, spores, paraphyses, anamorphs and chemistry (with either 'prasina 

unknown A' or 'B') as more typical forms (with coloured apothecia and more robust thallus). 

.


and I have seen intermediates on many occasions. The 'micrococca' forms are found in heavily 

shaded situations (e.g. tree trunks in dense conifer plantations, and trunks of understory trees or 

shrubs) and I consider their diminutive stature and lack of pigmentation to be a phenotypic 

response to low light intensities. The specimens described as Micarea polytrichi are similarly 

diminished forms, in this case overgrowing mosses (mainly Polytrichum spp.) by woodland 

roads. 

European populations of M. prasina have been found to be represented by three chemical 

races, each with one of three distinctive, but as yet unidentified, substances (see p. 87). During 

the course of the present study I have been tempted to recognize the chemical races as formal 

varieties. However I defer from doing so at this time because: (a) the chemical structure and 

biogenetic relationships between the three substances is not yet known; (b) the three chemical 

races do not consistently correlate to any clear differences in morphology; (c) the correlations 

between chemistry and distribution and (or) ecology are as yet uncertain. In the British Isles (at 

least) the race containing the 'unknown A' is by far the commonest and is a ubiquitous coloniser 

of trees and shrubs in all types of woodland (including young conifer plantations), occurring on 

tree boles, bark and lignum of fallen trees and stumps, and fallen sticks; it also occurs on shaded 

mossy turf on the ground and amongst rocks (usually with an east- or north-facing aspect) in 

maritime situations. The race containing 'unknown B' has a more restricted ecology and is 

mainly found in old woodland situations (including the native pine-woods), growing on the 

rather dry lignum or old bark of large stumps or fallen trunks; it has not been found on coastal 

turf or rocks. Specimens of this race tend to have a thick, pale grey (often abrading to greenish 

Map 18 Micarea prasina sensu lato • 1950 onwards O Before 1950


Map 19 Micarea prasina substance A # 1950 onwards O Before 1950 

white) thallus, but equivalent morphological forms containing 'unknown A' are sometimes 

encountered. Specimens containing 'unknown C are exceedingly rare and are known only from 

coastal turf in south-west England and the Channel Islands, and on argillaceous soil on a bank by 

a woodland path in south-eastern Schleswig-Holstein. The above ecological and phytogeographical 

tendencies require to be more thoroughly tested throughout the range of M. prasina, 

and it is possible that a good case for the taxonomic recognition of these races as varieties, 

subspecies, or even species could be made. 

The type material of Lecidea subviridescens contains 'unknown C and has spores that are at 

the high end of the overall size range for M. prasina s. ampl. (10-18x4-5-5 /xm) and some with 

two or three septa. However the very large 2-3-septate spores are always old with secondarily 

thickened walls, and similar spores are sometimes encountered in other Micarea species that 

have predominantly 1-septate spores. Furthermore, similarly large, 2-3-septate spores have 

been found in some coastal specimens containing 'unknown A' and in some inland, lignicolous 

specimens containing 'unknown B'. Although I regarded 'subviridescens'' as a separate species in 

the recent British checklist (Hawksworth etal., 1980: 62), I now believe it should be subsumed 

under M. prasina pending further studies as intimated in the preceding paragraph. 

Habitat and distribution (see also 'observations'): As an epiphyte on trees or shrubs M. 

prasina has been found on a wide range of phorophytes; in Britain these include Acer, Alnus, 

Betula, Castanea, Corylus, Fagus, Fraxinus, Metrosideros, Quercus, Salix, Sambucus, Sorbus, 

Ulmus, Calluna, Erica, Rhododendron, Vaccinium, Sarothamnus, Ulex, Abies, Larix, Picea^ 

Pinus, Pseudotsuga, and Juniperus. It occurs on the bark or lignum (old dried wounds, etc.) of


Map 20 Micarea prasina substance B # 1950 onwards O Before 1950 substance C ® Before 1950 I 

shaded trunks or main stems, but also on low (shaded) branches, fallen debris (especially in 

conifer plantations) and stumps. It is usually found in woodlands but sometimes occurs in 

sheltered niches in situations, e.g. bases of Ulex stems in more open, exposed hillside 

gorse-scrub. 

The phytosociological affinities of M. prasina are various and difficult to define. When on the 

bark of deciduous trees it sometimes occurs in shaded facies of communities belonging to the 

Lobarion pulmonariae, Parmelietum revolutae, and the Pseudevernietum furfuraceae. It is 

frequently found in small bark crevices amongst ± smooth areas of bark which are colonized by 

communities of the Graphidetum scriptae, Pyrenuletum nitidae, and Lecanoretum subfuscae. 

When occurring in such situations as the shaded trunks of young conifers or shrubs M. prasina 

often exists as ± pure stands; more detailed phytosociological studies may well lead to the 

formal syntaxonomic recognition of such communities. From amongst the British collections in 

E the associated species of M. prasina on the bark of deciduous trees include Arthonia spadicea, 

Bacidiavezdae, Chrysothrix candelaris, Cladonia coniocraea, Dimerella diluta, Evernia prunas- 

tri, Graphis elegans, Gyalideopsis anastomosans, Hypogymnia physodes, Lecidea icmalea, 

Lepraria incana, Ochrolechia androgyna, Opegrapha vulgata, Pachyphiale cornea, Parmelia 

glabratula, P. sulcata, Pertusaria multipuncta, Thelotrema lepadinum, Trapelia corticola, Dicranoweissia 

cirrata, Frullania spp., Hypnum cupressiforme, Lejeunea ulicina, and Metzgeria 

fareata. 

When on lignum its affinities are similarly difficult to define but it is often found in more 

sheltered or shaded (humid) niches adjacent to communities of the Calicietum abietini, 

Cladonietum coniocraeae, Lecideetum ostreatae, and Parmeliopsidetum ambiguae.


Species associated with M. prasina on the hgnum of fallen trunks and large stumps in the 

British Isles include Bryoria fuscescens, Chaenotheca ferruginea, Chaenothecopsis spp., Cladonia 

chlorophaea agg., C. macilenta, C. ochrochlora, Hypocenomyce scalaris, Hypogymnia 

physodes, Lecanactis abietina, Lecidea aeruginosa, L. icmalea, Micarea adnata, M. melaena, 

and Platismatia glauca. 

In coastal districts M. prasina is often found on plant debris, soil or moribund bryophytes in 

rock crevices or on ledges in sheltered gullies, and sometimes it occurs on the ground growing 

over plant debris or old Armeria tussocks, etc. Such habitats are usually sheltered and (or) with a 

north- to east-facing aspect. Associated lichens are few, but include Cladonia spp., Lecidea 

granulosa, L. icmalea, and Lepraria incana agg. Finds ofM. prasina growing directly on rock are 

very rare, but I have seen a few collections made from sandstone rocks in woodlands (from east 

Sussex and north-east Yorkshire). 

M. prasina is still to be found close to the centres of large conurbations and cities (e.g. Bristol, 

Edinburgh, London, and the West Yorkshire conurbation), and its persistence is probably due 

more to its ability to avoid, rather than tolerate, the direct effects of air pollutants such as 

sulphur dioxide and its derivatives. It naturally favours substrata with a low pH and is able to 

grow in very sheltered and shaded situations where more light demanding pollution resistant 

species (especially Lecanora conizaeoides) are at a competitive disadvantage. 

M. prasina s. ampl. is widely distributed in the British Isles and much of Europe. In 

Scandinavia it occurs northwards to at least c. 67°N. It may be rare in districts adjoining the 

Mediterranean Sea, but those areas are mostly poorly known lichenologically, and I have seen 

one collection from Toscana (northern Italy). M. prasina is present in Macaronesia (Azores and 

Canary Islands) and I have seen collections of it from several states in the U.S.A. (Georgia, 

Maryland, Massachusetts, Michigan, and Wisconsin). In addition I have seen several collections 

from South America and Australisia which come close to M. prasina, but these require more 

critical study and will be treated in a later publication. 

Exsiccata. Containing 'prasina unknown A': Arnold Lich. Exs. 1122, 1472 (BM ex K, M). Korber Lich 

Sel. Germ. 250 (L). Lojka Lich. Univ. 29 (BM ex K, M), 30 (BM ex K). Malme Lich. Suec. 23 (M, S) 

Rabenh. Lich. Eur. 676 (BM, BM ex K, M, WRSL). Rasanen Lich. Fenn. 651 (BM, BM ex K, M), 652 

(BM, BM ex K). Vezda Lich. Sel. 90 (BM, M), 1467 (BM). Zwackh. Lich. Exs. 593A (H-NYL 21601) 

Containing 'prasina unknown B': Arnold Lich. Exs. 280C (BM, GZU, M, WRSL). Cumm. Dec. N. Am 

Lich. I. 355 (BM, WIS). Hepp Flecht. Eur. 278 (BM, E, M, WRSL). Lojka Lich. Univ. 31 (BM ex K, M) 

Chemistry not tested: Arnold Lich. Exs. 279 (WRSL), 280A (BM, BM ex K, M), 280B (BM ex K, M) 

Arnold Lich. Mon. 243 (UPS), 245 (BM ex K, M). Hepp Zur. 224 (BERN). Kutak Lich. Bohem. 310 (O) 

Magnusson Lich. Sel. Scand. 134 (BM). Malme Lich. Suec. 24 (M, S). Rabenh. Lich. Eur. 733 (H) 

Rasanen Lich. Fenn. 653 (LD). Schaerer Lich. Helv. 196 p.p. (BM ex K, M). Vezda Lich. Sel. 1595 (BM 

GZU, M). Zahlbr. Lich. Rar. 175 (BM). Zwackh. Lich. Exs. 416 (UPS), 591A (H-NYL 21598), 591B 

(H-NYL 21594, 21599), 592A (UPS), 592B (M, UPS), 592C (UPS), 592D (M, UPS), 592E (UPS), 593B 

(H-NYL 21600), 593C (H-NYL 21595), 656 (UPS). 

35. Micarea pycnidiophora Coppins & P. James 

(Fig. 28: Map 21) 

in Lichenologist 11: 153 (1979). Type: England, South Hampshire, New Forest, near Cadnam, Shave 

Wood, 45 m, on bark of Fagus, 5 xi 1972, B. J. Coppins & F. Rose (E - holotype!; BM - isotype!). 

Thallus corticolous, often overgrowing moribund thaUi of bryophytes and other hchens; 

effuse, often wide-spreading, thin, uneven, composed of scattered to confluent areolae arising 

from a thin varnish-like prothallus. Areolae flattened to convex-hemispherical, grey-green or 

dull green, c. 40-100 îm diam; in section, without a distinct cortex or hyaline amorphous 

covering layer. Phycobiont micareoid, cells 4-7 /^m diam. 

Apothecia usually few or absent but sometimes abundant, immarginate, convexhemispherical 

to globose, often tuberculate, ivory-white to palUd, translucent when moist, 

surface matt, 0- 1-0-3 mm diam. Hymenium 35-50 /xm, hyaUne. Asci clavate, 30-35 x 10-12 /am. 

Spores ± straight and vertically aligned in the ascus (never tightly spiralled), shortly acicular.


Upper end usually broader and more obtuse than the lower end, usually slightly curved, 

3-7-septate, (14-)21-34x2-2-5(-2-7) (xm. Paraphyses numerous, branched and anastomosing, 

c. 1-1-5 ^tm, not swollen at apices. Hypothecium 20-80 ^tm tall, hyaline; hyphae interwoven, but 

becoming vertically orientated towards the hymenium, c. 1-1-5 /xm wide; ascogenous hyphae 

with swollen cells, c. 2-4 /xm wide. Excipulum poorly developed (c. 10 /am wide); hyphae 

radiating, branched and anastomosing, c. 1 /xm wide. 

Pycnidia always numerous and conspicuous, sessile or shortly stalked, whitish (concolorous 

with the apothecia), 100-300 /xm tall (including stalk) and 60-120 /xm diam; stalks (pycnidiophores) 

simple but occasionally clustered and appearing as if branched at the base; stalk 

tissue composed of interwoven hyphae c. 1-1-5 /am wide that ± separate in K. Conidiogenous 

cells ± cylindrical, 5-10x1-1-5 (xm. Conidia (mesoconidia) cylindrical, eguttulate, 3-8-6x1l-2(-l-5)/xm. 

acid. 

Chemistry: Apothecia, pycnidia and thallus (in section) K-, C+ red, PD-; t.l.c: gyrophoric 

Observations: Micarea pycnidiophora is characterized by its whitish apothecia and concol- 

orous, ± stalked pycnidia, shortly acicular spores, and C-l- red reactions (gyrophoric acid). It 

most closely resembles M. stipitata, which differs in having more elongate and often distinctly 

branched pycnidiophores, larger conidia, and C— reactions (gyrophoric acid absent). M. 

globulosella and M. synotheoides have similar acicular spores, but their apothecia are darkcoloured 

with an olivaceous (K-l- violet) pigment, and their pycnidia are inconspicuous and 

usually immersed in the thallus. Confusion could arise with Scoliciosporum pruinosum (P. 

Map 21 Micarea pycnidiophora k. + Micarea stipitata^


James) Vezda and 5". schadeanum (Erichsen) Vezda, but those species have narrower spores 

which are spiralled in the ascus and non-micareoid phycobionts (? Trebouxia); moreover, S. 

pruinosum has a granular epithecium (granules dissolving in K). Furthermore, these species 

have inconspicuous (? unknown) pycnidia, and do not contain gyrophoric acid. 

For illustrations of conidia and conidiogenous cells, and for further discussions, see Coppins 

& James (1979). 

Habitat and distribution: M. pycnidiophora is a species of old woodlands, in damp, sheltered, 

and rather shaded situations. It is generally found on areas of ± smooth-bark on the boles of old 

trees, especially Fagus and Ilex, but also Alnus, Quercus, and Rhododendron. Although 

characteristic of acid bark it has not yet been recorded on conifers. 

Its known centre of distribution is the New Forest of Hampshire in southern England, where it 

is locally frequent. Its other British localities are Sheffield Park in Sussex (on Rhododendron 

with Gyalideopsis anastomosans) and the Limb Valley, Yorkshire (its most northerly locality; at 

base of Fraxinus). The latter site is an ancient oak-wood just outside the large industrial city of 

Sheffield, but M. pycnidiophora occurs in an extremely sheltered situation, where it presumably 

avoids the effects of the high levels of air pollution prevalent in that area. M. pycnidiophora is 

little known outside of Britain, but is reported from Bretagne, France (Coppins, 1971, as 

'Bacidia sp. 2') and the Canary Islands. 

36. Micarea rhabdogena (Norman) Hedl. 

(Fig. 29A) 

in Bih. K. svenska VetenskAkad. Handl. Ill, 18 (3): 75, 85 (1892). - Biatora (Lecidea) rhabdogena 

Norman in Ofvers. K. VetenskAkad. Fork. 11: 803 (1870). - Lecidea rhabdogena (Norman) Th. Fr., 

Lich. Scand. 2: 473 (1874). Type: Norway, Nordland, Maalselven, Skjeggenaes, J. M. Norman (O - 

lectotype!; isolectotypes: BM!, O!, S!). 

Lecidea glomerellaf. ecrustacea Nyl. ex Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 457 (1934). - Lecidea 

glomerella f. ecrustacea Nyl. in Elfving in Medd. Soc. Fauna Fl. fenn. 2: 168 (1878); nom. nudum (Art. 

32). Type: USSR, Karelskaya ASSR, Karelia olonetsensis, Gorki near Svir, F. Elfving (H-NYL 19120- 

holotype!). 

Thallus inapparent, endoxylic; hyphal walls often olivaceous, K+ violet. Phycobiont probably 

micareoid, cells c. 5-7 ixm diam. 

Apothecia numerous, immarginate, convex-hemispherical or tuberculate, black, matt, 0-1- 

0-3 mm diam, or up to 0-4 mm diam when tuberculate. Hymenium 23-30 /xm tall; upper part 

(epithecium) fuscous-brown, HNO3— , K- but pigment dissolving and fading into solution; 

remaining (lower) part dilute yellowish brown; hymenium sometimes tinged violet in K (evident 

after brown pigment has dissolved). Asci clavate, 19-28x7-9 /am. Spores oblong-ellipsoid, 

ovoid-oblong or bacilliform, simple or rarely a few 1-septate, 6-9xl-5-2-3 /am. Paraphyses 

numerous, hyaline throughout, c. 1 /xm wide in mid-hymenium but often widening above to T5 

/am; sparingly branched and sometimes anastomosing, but becoming richly branched above 

where their entangled apices, together with the brown pigment, form a ± well delimited 

epithecium. Hypothecium c. 70-160 /am tall, pale, dilute yellowish brown, K— , HNO3— 

Excipulum reflexed, dilute to dark fuscous-brown, K— but partly dissolving into solution; 

hyphae radiating, branched and anastomosing, c. 0-7-OT /am wide, sometimes widening to 1-5 

/am towards the outer edge. 

Pycnidia of two types: (a) partly immersed to ± sessile, black, matt, 40-80 /am diam; wall 

olivaceous, K-l- violet; conidia (mesoconidia) oblong-ellipsoid, oblong-ovate or obovate, c. 

3-5-4-7X1-4—T8 /am; (b) immersed between surface wood fibres, black, c. 4 /am diam; wall 

olivaceous, K-l- violet; conidia (microconidia) narrowly cylindrical, 4-4-5 xO-7-0-9 /am. 

Chemistry: Sections of apothecia C— ; material insufficient for analysis by t.l.c. 

Observations: M. rhabdogena resembles M. elachista in apothecial pigmentation and anatomy, 

but differs in its endoxylic thallus, smaller and mostly simple spores, and smaller, matt 

black pycnidia. M. rhabdogena is similar in appearance and spore characteristics to M. misella. 

.


but the latter has an oHvaceous, K+ violet hymenium (fuscous pigment lacking) and its 

mesoconidia are borne in stalked pycnidia. Diminutive, endoxylic forms of M. denigrata can be 

distinguished by the absence of a distinct (fuscous) epithecium, hymenium reacting K+ violet, 

C+ violet (due to olivaceous pigment) and, usually, C+ orange-red (gyrophoric acid). The 

mesoconidial states of M. denigrata and M. rhabdogena are very similar, but M. denigrata has 

longer (4-5-7-5 /xm) microconidia, and sometimes has curved macroconidia; macroconidia are 

apparently not produced by M. rhabdogena or the related M. elachista. 

Habitat and distribution: M. rhabdogena appears to be a rare or much overlooked, exclusively 

lignicolous species, known so far only from north Norway, mid-Sweden and Karelia. All 

collections are on rather hard conifer lignum, and associated species include Calicium trabinellum, 

Cetraria pinastri, Mycocalicium subtile, Parmeliopsis ambigua, P. hyperopia, Xylographa 

abietina, andX vitiligo. 

Exsiccata: Malme Lick. Suec. 20 (M, S). 

37. Micarea stipitata Coppins & P. James 

(Fig. 29B;Map21) 

in Lichenologist 11: 156 (1979). Type: Scotland, Argyll, Loch Creran, Glasdrum National Nature 

Reserve, on Betula, 27 v 1976, L. Tibell & Coppins 2357 (E - holotype!; isotypes: BM!, H!, UPS, hb 

Poeh!,hbVezda!). 

Thallus and apothecia: more or less identical in appearance and internal anatomy to those of 

M. pycnidiophora (q.v.). 

Pycnidia always present numerous and conspicuous, borne on distinct whitish stalks (pycnidiophores) 

which may be simple, bifurcate or to 5-branched, mainly 400-800 /xm tall and 

60-100(-150) fxm diam, often with small irregular clusters of ± superficially encapsulated algae 

(small granular-areolae). Stalk tissue composed of interwoven hyphae, c. 1-1-5 /xm wide, which 

± separate in K. Pycnidia innate in the apex or apices of pycnidiophores, ± globose or doliiform, 

c. 60-90 fxm diam. Conidiogenous cells ± cylindrical, 7-11x1-4-2 /xm. Conidia (mesoconidia) 

cylindrical or narrowly ellipsoid, eguttulate, 6-8x1-1-8 fxm. 

Chemistry: All parts K- , C- , KC- 

, PD- 

; no substances detected by t. I.e. 

Observations: See under the related species M. pycnidiophora for the differences between 

them and for comparisons with other similar species. 

Habitat and distribution: M. stipitata is characteristic of acid, often leached, bark (or 

overgrowing bryophytes thereon) on trees in undisturbed woodlands in areas with a high 

rainfall. It occurs in communities referable (or closely akin) to the Parmelietum laevigatae on the 

trunks of Betula, Quercus, and, less frequently, Alnus, Pinus, Abies, and Pseudotsuga. It has 

once been found on mossy rocks. 

The British distribution of M. stipitata is a distinctly more northern and western, as well as 

more oceanic, than that of M. pycnidiophora (see Map 21). It has not yet been found in 

south-west England but presumably occurs there. Although known from the Azores and the 

Canary Islands it has not been found elsewhere in Europe, but should be looked for in the 

hyperoceanic woodlands of, for example, south-west Norway and France (Bretagne). 

Additional information: for additional illustrations and some further discussion see Coppins & 

James (1979). 

38. Micarea subleprosula (Vezda) Vezda 

(Figs 30, 53-54; Map 25) 

in Vezda & V. Wirth in Folia geobot. phytotax, Praha 11: 101 (1976). - Bacidia subleprosula Vezda in 

Preslia 33: 366 (1961). Type: Czechoslovakia, Bohemia, Sudety, Krkonose, Mumlava valley near 

Harrachov, 900 m, over decaying mosses on granitic rocks, 1960, A. Vezda (E - isotype! [t.l.c: 

alectorialicacid]). 

^


Thallus: As for M. leprosula {q.v.) but with a different chemistry (see below). 

Apothecia sometimes (? often) absent, immarginate, ± globose and constricted at the base, 

sometimes irregularly aggregated or tuberculate, grey-brown, brownish black, or black and 

often with bluish tinge, 0-2-0-8 mm diam. Hymenium c. 65-90 /xm tall, ± hyaline but upper part 

oHvaceous or dilute reddish brown, K— , HNO3-I- reddish. Asci clavate, c. 60-70x19-22 /am. 

Spores fusiform, often slightly curved, 3-7(-9)-septate, (35-)40-50(-60) /xm. Paraphyses 

numerous, branched and often anastomosing, 1-1-5 /itm wide; apices often more richly 

branched, not or only slightly incrassate (to 1-8 /xm). Hypotheciumc. 120-200 /xm tall, ± hyahne 

or dull straw, but often mottled dilute fuscous-brown (K-) in the upper part; hyphae 

interwoven, but becoming outwardly orientated towards the hymenium and excipulum, c. 

0-8-1 -5 /xm; ascogenous hyphae with swollen cells, c. 2-6 fxm wide. Excipulum distinct in young 

apothecia, but reflexed and obscured in old apothecia, hyaline or dilute fuscous-brown in part; 

hyphae radiating, branched and anastomosing, c. 1-1-5 jxm wide. 


Chemistry: Thallus K— , C+ 

acid plus two unidentified accessory substances. 

red, PD-I- yellow; sections of apothecia C— ; t.l.c: alectorialic 

Observations: Micarea subleprosula is closely allied to M. leprosula and sterile thalli of the two 

species can only be distinguished by their reactions with PD, or by t.l.c. See under M. leprosula 

for further discussion. 

For additional illustrations of A/, subleprosula see Vezda (1961). 

Habitat and distribution: M. subleprosula grows over mosses on rocks in much the same 

habitats in which M. leprosula is mostly commonly found. It was originally described from the 

Sudety mountains of Czechoslovakia, where it occurred at an altitude of 900 m, and I have seen 

only two additional specimens: from Sweden (Varmland) at 200 m; and Wales (Snowdonia) at c. 

760 m. It may have been overlooked for M. leprosula. However, I have tested numerous 

individual thalli personally gathered in the field (mainly in Scotland) during the last six years, 

and all of these were Pd+ red and referable to M. leprosula. 

39. Micarea subnigrata (Nyl.) Coppins & Kilias 

(Figs. 31A, 50: Map 22) 

in Kilias in Herzogia 5: 391 (1980). - Lecidea subnigrata Nyl. in Flora, Jena 49: 370 (1866). - Lecidea 

denigrata* [subsp.] subnigrata (Nyl.) Crombie, Lich. Brit.: 70 (1870). - Catillaria subnigrata (Nyl.) 

Herre in Proc. Wash. Acad. Sci. 12: 94 (1910). Type: Wales, Merioneth, Cader Idris, 1866, W. A. 

Leighton (H-NYL 19136 -lectotype!, sel. Kilias (/oc. dr.); BMexK-isolectotype!). 

Lecidea confusula Nyl. in Flora, Jena 55: 360 (1872). - Micarea confusuta (Nyl.) Hedl. in Bih. K. svenska 

VetenskAkad. Handl. Ill, 18 (3): 76, 86 (1892). Type: Scotland, East Perthshire, Blair Atholl, Craig 

Tulloch, on mica-schist, 1871, Crombie (H-NYL 20191 -lectotype!; BM-isolectotype!). 

Thallus effuse, widespreading or as small patches amongst other lichens, of scattered to 

confluent or clustered, irregularly convex to subglobose areolae. Areolae 0-08-0-45 mm diam 

(usually at their largest when adjoining apothecia), dark grey-brown, matt or slightly glossy; in 

section with a hyaline amorphous covering layer c. 3-10 jxm thick, and outermost hyphae often 

pale brown (K-); a white medulla often developed in large areolae. Phycobiont mvc^nQOid, cells 

c. 4-7 /xm diam. 

Apothecia usually numerous, scattered to crowded and confluent, immarginate, at first adnate 

and slightly convex, soon becoming convex-hemispherical and occasionally tuberculate, dark 

brown (reddish brown when wet) to brownish black, matt or slightly glossy (especially when 

young), 0-2-0-6 mm diam, or up to c. 1 mm diam when tuberculate. Hymenium 35-38 /am tall, 

hyaline but upper c. 10-12 fxm (epithecium) fuscous brown, K-, HNO3-. Asci clavate 

33-35x10-14 /xm. Spores ellipsoid (0-)l -septate, 8-12x4-5 /am. Paraphyses numerous, 

branched and anastomosing, (l-)l-3-l-7 /xm wide, sometimes, gradually widening to 2-5 fxm 

towards apices; hyaline throughout (epithecial pigment confined to gel-matrix). Hypothecium


45-160 ixm tall, hyaline; hyphae interwoven, some becoming vertically orientated towards the 

hymenium, c. 1-2-1 -8 /xm wide, intermixed with ± inflated, short-celled ascogenous hyphae c. 

2-5-5 fjcm wide. Excipulum well developed, 25-45 yam wide, dilute fuscous brown; hyphae 

radiating, branched and anastomosing, c. 1-3-1-7 jxm wide. 

Pycnidia usually present, immersed within areolae with ostioles visible (x50) as minute pores 

surrounded by a thin, often slightly raised, dark brown rim; in section with a simple circular or 

flask-shaped cavity, walls hyaline, or pale brown around the ostiole; of two types: (a) c. 80-200 

/xm diam; conidiogenous cells ampulliform to subcylindrical, some with 1-2 percurrent proliferations, 

4-10 X 1-5-2-5 /xm, sometimes with swollen base to 4 /xm wide; conida (macroconidia) 

helicoid with 2-3 spirals, often appearing 3-5-septate, overall size 7-10x5-6 /xm, filaments 

1-8-2 /xm wide (b) c. 40-100 /xm diam; conidiogenous cells ± cylindrical or ampulliform, 

4-lOx 1-1 -4 /xm, sometimes with swollen base to 3 /xm wide; conidia {microconidid) cylindrical, 

4-6-6x0-8-1 /xm. 

Chemistry: Thallus K— , C-, PD — ; sections of apothecia C-; t.l.c: no substances. 

Observations: Micarea subnigrata is characterized by its dark grey-brown, verrucose-areolate 

thallus, dark brown apothecia, fuscous brown epithecium and excipulum, colourless hypothecium, 

and 0-1-septate, eUipsoid spores. However, its most outstanding and characteristic 

feature is the helicoid shape of its macroconidia (Fig. 50A); such conidia are unique in the genus 

and apparently so in all pycnidial fungi, whether lichenized or not. In general appearance, 

habitat and shape and septation of spores M. subnigrata could be confused with M. intrusa, but 

Map 22 Micarea intrusa A 1950 onwards A Before 1950 + Micarea subnigrata • 1950 onwards O Before 

1950 

i


that species has a green epithecium and a large-celled, non-micareoid phycobiont. The little 

known M. curvata may be closely related to M. subnigrata, but can be distinguished by the 

fabiform or strongly curved spores and C+ red reaction of apothecial sections. With its external 

and internal fuscous brown colourations M. subnigrata could be mistaken for a Fuscidea, but 

species of that genus have broader (c. 1 •7-2-7 /xm) mostly unbranched paraphyses (the apical 

cell of each being clavate or capitate and often provided with a dark brown 'apical cap'), asci with 

a thick, strongly amyloid apical wall, and probably referable to the Teloschistes-type (Honegger, 

1978), and a protococcoid phycobiont, with thick-walled cells, c. 10-14 îm diam. 

For additional illustrations of M. subnigrata see Kilias (1981: 391, 445). 

Habitat and distribution: M. subnigrata is confined to hard siliceous rocks in rather exposed 

situations. The communities in which it occurs are attributable to the Lecideion tumidae 

alliance, especially the Lecideetum lithophilae association. The collection {Coppins 8417) from 

Glentrool in Kirkcudbrightshire was made from the south-facing side of a doleritic boulder on a 

south-facing slope, at an altitude of 135 m; association species were: Cladonia subcervicornis, 

Huilia albocaerulescens, H. tuberculosa, Lecanora badia, Lecidea pycnocarpa, Lepraria neglec- 

ta, Rhizocarpon obscuratum, R. oederi, Stereocaulon vesuvianum, Trapelia involuta, T. aff. 

obtegens, and Andraea sp. From other British collections the following can be added: Candela- 

riella vitellina, Catillaria chalybeia, Lecanora intricata, L. polytropa, Lecidea fuliginosa, L. 

lactea, Lecidella scabra, and Parmelia verruculifera. For information on associated species in its 

Norwegian locality see M. intrusa (Table 5). 

M. subnigrata is known from scattered localities in upland districts in the Scottish highlands, 

Galloway, and Wales; recent field studies indicate that it may be far more common in these and 

similar areas (e.g. Lake District and Dartmoor) than present records (Map 22) suggest. Outside 

Britain it is known only from coastal Norway (Hordaland) and south-west Sweden (Halland). 

40. Micarea subviolascens (Magnusson) Coppins, comb. nov. 

(Fig. 31B) 

Lecidea subviolascens Magnusson in Blyttia 7: 30 (1949). Type: Norway, Hordaland, Granvin, Steinsaethorgen, 

on easily weathered schist in an open windy place of a small hill, 780 m, ix 1944, J. J. Havaas, 

Lich. Exs. Norv. 694 (BG-lectotype!). Topotype material collected in 1949distributed in Havaas, L/c/z. 

Exs. Norv. 710 (BG!) and Lich. Norv. Occid. 269 (BG!). 

Lecidea assimilata f. aberrans Th. Fr., Lich. Scand. 2: 523 (1874). Type: Norway, Troms, Troms0, 

Fl0jfjellet, 1868, Th. M. Fries (UPS-holotype!). 

Lecidea assimilata var. hardangeriana Vainio in Acta Soc. Fauna. Fl. fenn. 57 (2): 374 (1934). - Lecidea 

assimilata var. hardangeriana Vainio in Havaas in Bergens Mus. Arb. 1909 (1): 29 (1909); nom. nudum 

(Art. 32). Type: Norway, Hordaland, Granvin, Sm0reggen, 650 m, 22 viii 1902, J. J. Havaas, Lich. Exs. 

Norv. 139 (BG-lectotype! [t.l.c.: no substances]; isolectotypes: BG!, H!). 

Thallus effuse, saxicolous, composed of ± confluent, white to pale brown, convex, verrucoseareolae 

c. 0'l-0-5(-0-8) mm diam; with age the crust may thicken (to c. 1 mm) and become 

cracked and divided into 'islands' c. 1-2 /xm wide. Areolae in section, sometimes with a hyaline 

amorphous covering layer c. 5-7 /xm thick, upper c. 20 /xm of areolae often tinged dilute 

olivaceous and Kf+ violet (pigment confined to the weak gel-matrix). Thallus hyphae 1-7-2-5 

(-3) /xm wide. Phycobiont micareoid, cells 4-7 /xm diam. Cephalodia (?): spaces between 

areolae often filled by black, loose clusters of Stigonema. 

Apothecia numerous, immarginate, adnate, convex to subglobose, black, matt, 0-3-0-6(-0-8) 

mm diam, often confluent, or forming tuberculate clusters up to 1-2 /xm diam. Hymenium 40-55 

/xm tall, dark green (K+ violet, HNO3+ purple-red) above, and below in vertical streaks, 

otherwise dilute greenish or ± hyaline. Asci clavate, 38-48x10-15 /xm. Spores ellipsoid, 

ovoid-ellipsoid or oblong-ellipsoid, simple, 9-16(-17)x4-5 /xm. Paraphyses numerous, simple 

below, but in upper part often forked or with short lateral branches, l-l-8(-2) /xm wide, 

sometimes widening above to 3 /xm; apical walls hyaline although surrounded by densely 

pigmented gel-matrix. Hypothecium 150-300 /xm tall, dark purple-brown, K4- purple intensifying, 

or upperpart K-l- dark green; all parts HNO3+ purple-red; hyphae interwoven, but ±


vertically arranged in upper part, 1 •5-2-5 îm wide, embedded in densely pigmented gel-matrix; 

ascogenous hyphae c. 2-5-5 /xm wide. Excipulum ± distinct in young apothecia, but soon 

reflexed, dark purple-brown (K-l- dark green) within, changing to green (K-l- green intensifying) 

towards the outer edge; hyphae radiating, branched, c. 1-5-2 fxm wide. 

Pycnidia: Not found. 

Chemistry: Thallus K— , C— , KC— , PD— ; t.l.c: no substances. 

Observations: In his protologue, Magnusson allied his new species with the Lecidea [Micarea] 

sylvicola group. However, Th. M. Fries, 75 years earlier, was probably nearer the truth when he 

described material of M. subviolascens as a form {'aberrans') of Lecidea [Micarea] assimilata. In 

my opinion M. subviolascens is very closely related to M. assimilata, and the reader is referred to 

the account of that species for further discussions. 

Habitat and distribution: M. subviolascens appears to have a very restricted distribution, being 

known only from the provinces of Troms and Hordaland in Norway. It is the only known 

saxicolous member of the M. assimilata group, and occurs on acidic rocks (schists and gneiss) in 

rather open situations. In the Hordaland localities it was collected at altitudes of 650 m and 795 

m. The labels accompanying the specimens provide little extra ecological information, and the 

specimens have few associated species, although Pyrenopsis pulvinata and Rhizocarpon obscuratum 

have been noted. 

Exsiccata: Havaas Lich. Exs. Norv. 139 (BG, H), 694 (BG), 710 (BG). Havaas Lich. Norv. Occid. 269 

(BG). 

41. Micarea sylvicola (Flotow) Vezda & V. Wirth 

(Figs31C,51A;Map23) 

in Folia geobot. phytotax, Praha 11: 99 (1976). - Lecidea sylvicola Flotow, Lich. Schles. 171 (1829). 

Type: Poland, Schlesien, Flotow, Lich. Exs. 171A (UPS - lectotype!, sel. Hertel (1975: 74); WRSL- 


Lecidea aggerata Mudd, Man. Br. Lich.: 208 (1861). Type: England, Yorkshire, Battersby, Mudd, Lich. 

Brit. 175 (BM - lectotype! ; isolectotypes: E!, H-NYL 14016!, M!, MANCH!). 

Lecidea incincta Nyl., Lich. Scand. : 231 (1861). Type: Finland, Satakunta, Kallfjard [Ahlainen: Kellahti], 

1859, /i. /. Malmgren (H - holotype!). 

Biatora smaragdina ArnoXd'm Verh. zool. bot. Ges. Wien 19: 613 (1869). Type: Italy, Trentino-Alto Adige, 

'Melaphyr in Walde unterhalb Bad Razzes am Schiern [Monte Pez]', vii 1867, Arnold (M - holotype!). 

Lecidea hellbomii Lahm in Flora, Jena 53: 177 (1870). Type: Sweden, Narke, Vredstorp, Urby, on granitic 

rock, 1869, P. J. Hellbom (M- lectotype!). 

Lecidea sylvicola f. sublivida Vainio in Medd. Soc. Fauna Fl. fenn. 10: 104 (1883). - Lecidea sylvicola var. 

sublivida (Vainio) Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 298 (1934). Type: Finland, Ostrobottnia 

kajanensis, Kuhmo,Kylmala, 1877, E. A. Vainio (TUR-VAINIO 25226 -holotype!). 

Lecidea hypocyanea Vainio in Acta Soc. Fauna Fl.fenn. 57 (2): 300 (1934), non Stirton (1879). - Lecidea 

vainioi Magnusson in Blyttia 7: 31 (1949); nom. nov. Type: Finland, Regio aboensis, Turku, Hirvensalo, 

10 vii 1924, E. A. Vainio (TUR-VAINIO 33172 -lectotype!, sel. Hertel (1975: 74)). 

Lecidea sylvicola \ar.flotowii Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 297 (1934); nom. inval. (Art. 26). 

Thallus effuse, thin and ± smooth or weakly rimose to rather thick, uneven and coarsely 

rimose, or sometimes with irregularly granular or verrucose areolae up to 0-3 mm diam, pale 

buff, pale to dark grey or bluish grey; in section without a cortex or hyahne covering layer, but 

walls of outermost hyphae sometimes greenish, K— , HNO^-f- purple-red. Phycobiont not 

micareoid; cells thin-walled, ± globose and c. 5-12 /xm diam, or ellipsoid and up to 15x 10 fxm. 

Apothecia numerous, convex-hemispherical from the start, often becoming ± globose or 

tuberculate, black and often with bluish tinge, sometimes dark blue-grey (deep shade forms), 

0-2-0-5 diam, or to 1-2 mm when tuberculate. Hymenium 40-60(-70) /xm tall, dilute bright or 

sordid aeruginose, but often darkish aeruginose in upper and lower parts, and in vertical streaks 

(due to presence of stout pigmented paraphyses), K— or -I- aeruginose intensifying, HNO3-I- 

purple-red. Asci cylindrical-clavate, c. 35-45x8-12 /xm. Spores ellipsoid or ovoid, simple (an 

occasional 1-septate spore seen in a few collections), (6-)7-10x(2-5-)3-4-5 /am. Paraphyses


rather scanty, of two types: p.p. evenly distributed, irregularly flexuose, simple or sparingly 

branched, often anastomosing, hyaline, thin, 0-7-1 /xm wide, sometimes widening above to 1-8 

fim; p.p. fewer in number, as scattered individuals or in small fascicles, straight, simple or 

occasionally forked above , stout, coated ± throughout (but most intensely so around the apices) 

by dark green pigment and appearing c. 2-3 /xm wide, apices sometimes incrassate and up to 4 

/Lim wide. Hypothecium 90-200 /xm tall, dark aeruginose or olive-black, K- or + green 

intensifying; lower ('core') part, or rarely the entire hypothecium, dark purplish brown, K-lpurple 

intensifying; all pigments HNO3+ purple-red; hyphae coated with dense dark green 

pigment and 2-3 /xm wide (overall), interwoven but becoming vertically orientated towards the 

hymenium and some continuing into it as stout, pigmented paraphyses; ascogenous hyphae 

similarly pigmented, with swollen cells c. 3-5 /xm wide. Excipulum not evident, even in sections 

of young apothecia. 

Pycnidia usually present and numerous, immersed, black, 40-200 /xm diam; in section with a 

single circular or ovate locule but often becoming internally convoluted with up to 6 locules, wall 

dark green K- , HNOs-f- purple-red in upper part, turning paler below and dilute brownish or ± 

hyaline at the base; conidiogenous cells irregularly subcylindrical, 5-10 x 1-2-1 -5 /xm, often with 

one or two percurrent proliferations; conidia (mesoconidia) ± cylindrical or sometimes oblongobovoid, 

sometimes biguttulate and often ± constricted in the middle, 3-8-6(-6-6)xl-l-7(-2) 

/xm. 

Chemistry: Thallus K-, C-, KC-, PD-; sections of apothecia C-; t.l.c: no substances. 

Observations: Micarea sylvicola is characterized by its convex-globose, often tuberculate, 

black apothecia, green or blue-green hymenium, very dark green-black or brown-black 

hypothecium (appearing green, or rarely purplish, in K), and ellipsoid to ovoid, simple spores. 

The variation in hypothecial pigmentation is similar to that found in such species as M. 

assimilata, M. crassipes, and M. melaena. M. tuberculata is rather similar to M. sylvicola but has 

generally smaller apothecia, a more shallow hymenium, narrower, oblong-ovoid spores that are 

often 1-septate, and shorter conidia. Forms of M. bauschiana with greenish pigment in the 

hypothecium have been confused with M. sylvicola, however the pigment is always dilute and 

confined to the gel-matrix such that the hyphae appear hyaline in K (at x 1000) . The thallus of M. 

sylvicola is often provided with a blue-grey tinge due to green-pigmented hyphae near the 

surface; such coloration has not been noticed with the thalli of M. bauschiana and M. 

tuberculata. Another species often confused with M. sylvicola is Lecidea erratica Korber, which 

can be distinguished by its thinly marginate young apothecia, and distinct excipulum when 

observed in section. 

Habitat and distribution: M. sylvicola is mainly found on dry shaded rocks in the communities 

belonging to Micareetum sylvicolae, but it sometimes occurs in periodically wetter situations on 

rock faces, upper sides of boulders, and loose stones in woodlands where it may be associated 

with such species as Baeomyces rufus, Cystocoleus ebeneus, Fuscidea recensa, Huilia tuberculosa, 

Parmelia saxatilis, Rhizocarpon hochstetteri, R. obscuratum, R. oederi, Scoliciosporum 

umbrinum, Trapelia involuta, and in one instance (Coed Hafod, Denbigh) Bacidia vezdae. In 

addition, it is sometimes found on old fence posts in upland districts. 

In the British Isles M. sylvicola is more confined to upland districts and is less common than M. 

bauschiana. It seems to be particularly prevalent in Wales and south-east Scotland, and 

curiously rare in western Scotland and Ireland. Further field-work is required to establish 

whether or nor its apparent scarcity in these latter areas is real or merely an artefact resulting 

from uneven recording. However, my field observations in western Scotland suggest that its 

niche in the Micareetum sylvicolae is there occupied by M. lutulata. M. sylvicola is widely 

distributed in Scandinavia, but rarely found from north of about latitude 67°N. Elsewhere in 

Europe I have seen specimens from Germany (Hessen and Baden-Wiirttemberg), eastern 

Austria, northern Italy, south-west Poland and the mountain regions of Czechoslovakia. From 

outside Europe I can confirm its presence in north-eastern North America (New York and 

Newfoundland).


Map 23 Micarea sylvicola # 1950 onwards O Before 1950 

Exsiccata: Arnold Lich. Exs. 409A (BM ex K, M, MANCH, WRSL), 409Bp./7. (BM ex K, M). Flotow 

Lich. exs. 171A (WRSL). Johnson Lich. Herb. 434 p.p. (BM). Kav. & Hilitzer Crypt. Cech. 269 (UPS). 

Korber Lich. Sel. Germ. 75 (M). Larb. Lich. Caesar. Sarg. 84 (ABD, BM, MANCH). Larb. Lich. Herb. 

304, 305 p.p. (BM). Malme Lich. Suec. 199 (M, S). Mudd Lich. Brit. 175 (BM, E, M, MANCH). Norrl. & 

Nyl. Herb. Lich. Fenn. 763 (BM, H, TUR, WIS), 764 (BM, H, WIS). Rasanen Lich. Fenn. 612 p.p. (M). 

Suza Lich. Bohem. 131 (M). Vezda Lich. Sel. 957, 1341 (BM). Zwackh Lich. Exs. 535, 596 (M), 597 

(UPS), 780 (M), 919 (BM, M). 

42. Micarea synotheoides (Nyl.) Coppins 

(Figs . 3 1 D-E , 43D-E ; Map 24) 

in Topham & Walker in Lichenologist 14: 67 (1982) - . Lecidea synotheoides Nyl . , Lich. Jap. : 63 ( 1 890) . - 

Catillaria synotheoides (Nyl.) Zahlbr., Cat. lich. univ. 4: 78 (1926). Type: Japan, 'Itjgome' (or 

'Itchigome'), on lignum, 1879, E. Almquist (H-NYL 19101 - lectotype!; S - isolectotype!). 

Thallus effuse, of scattered to coherent, or clustered, granular areolae. Areolae c. 20-70 /am 

diam, dull grey-green (never whitish) to dark olivaceous or blackish, appearing ± gelatinous 

when wet, usually ± discrete but often coalescing to form a thin uneven crust; a thin white 

prothallus sometimes visible between scattered areolae. Areolae in section without an amorphous 

covering layer; outer hyphae often with olivaceous or brownish, K-l- violet walls. 

Phycobiont micareoid, cells c. A-1 fxva. 

Apothecia immarginate, convex-hemispherical to ± globose, often tuberculate, grey-black or 

brown-black, matt, sometimes grey (shade forms), 0- 1-0-3 mm diam, or up to 0-5 mm diam


when tuberculate. Hymenium 35-45 /xm tall, dilute olivaceous or dilute olive-brown, K+ violet: 

pigment confined to gel-matrix. Asci clavate, 30-40x9-5-12 jxm. Spores ± acicular or rodshaped, 

curved or ± straight, l-7(-ll)-septate, 14^35(-43)xl-8-2-5(-3) y,m. Paraphyses 

numerous, branched, sometimes anastomosing, 0-8-1 /xm wide, sometimes widening to 1-5 /am 

towards their apices; apical walls hyaline. Hypothecium 60-100 jxm tall, hyaline, or dilute 

olivaceous and then Kf+ violet; hyphae hyaline, 1-2 îm wide, interwoven or some ± vertically 

orientated in the upper part; intermixed with short-celled ascogenous hyphae, c. 2-4 îm wide. 

Excipulum indistinct, evident in sections as a narrow, reflexed, non-amyloid lateral border to 

the hymenium, hyaline or pale olivaceous (then K+ violet), varying from paler to darker than 

the hymenium; hyphae hyahne, radiating branched and anastomosing, c. 1 /xm wide. 

Pycnidia frequent but inconspicuous, immersed within areolae, or emergent to sessile, 

whitish to grey-black; walls dull olivaceous (pigment often more intense around the ostioles), 

K+ violet. Pycnidia of two types: (a) 60-120 /am diam, immersed, or emergent with gaping 

ostioles; conidia {mesoconidia) ± cylindrical to fusiform, 4-5-6xl-2-l-5 /xm; (b) 30-40 /Ltm 

diam, immersed to sessile, but ostioles not gaping; conidia (microconidia) short-cylindrical, 

3-8-4-8x0-8-l/xm. 

Chemistry: Thallus C— , K— , PD— ; sections of thallus and apothecia C- (but olivaceous parts 

C-l- violet due to pigment); t.l.c: no substances detected. 

Observations: Micarea synotheoides is characterized by its ± acicular or rod-shaped spores, 

K-l- violet hymenium, and olivaceous thallus which is j. gelaimous when wet. M. globulosella is 

very similar, but differs in having a thallus of whitish or grey, somewhat larger areolae 

containing gyrophoric acid. M. nitschkeana differs in the same ways, and in addition has shorter 

(rarely over 17 jxm) spores and longer microconidia. M. globulosella and M. synotheoides have 

been confused with Bacidia beckhausii (see p. 196) which has similar spores, pigmentation and 

paraphyses, but can be distinguished by its large-celled (c. 8-14 fxm diam) phycobiont, 

excipulum of coherent (in K) hyphae, smaller conidia (one type only, c. 2-8-3-5X 1-1-4 /xm), 

often pruinose apothecia, and usual occurrence on less acidic bark (especially of Acer, Fraxinus, 

Populus, and Ulmus), although it does occasionally occur on Quercus trunks and more rarely on 

lignum. 

Of the four Japanese specimens, two are on bark and two (type material) are on lignum. They 

all have rather siiprt spores (14-26 /xm), but appear to agree in most other respects with 

collections from Britain and the Atlantic islands. Unfortunately I was unable to find mesoconi- 

dia in the Japanese specimens. For the time being, I consider the Japanese and Atlantic 

populations as conspecific, especially as the British collections exhibit considerable variation in 

spore septation and length. Further collections from Japan are required to finally establish the 

relationship between the two populations. 

Habitat and distribution: M. synotheoides is an oceanic species mostly found in wooded valleys 

on bark or over bryophytes, on the trunks of, for example, Alnus, Betula, Quercus, Larix, 

Pinus, and Pseudotsuga, in communities in, or related to, the Parmelion laevigatae alliance. 

British collections include the following associated lichens: Bryoria fuscescens, Chrysothrix 

candelaris, Cladonia coniocraea, C. squamosa, Haematomma elatinum, Hypogymnia physodes, 

Lecanactis abietina, Lecidea icmalea, Micarea alabastrites, M. cinerea, M. peliocarpa, M. 

stipitata, Mycoblastus sterilis, Ochrolechia androgyna, Parmelia saxatilis, Pertusaria amara, 

Platismatia glauca, Sphaerophorus globosus, Trapelia corticola in ed., and Usnea subftoridana. 

The single British collection on lignum {Coppins 2299) was on a rotting trunk of Pinus. In the 

Canary Islands it occurs on Erica arborea in a photophobus and hydrophilic community 

composed of crustose lichens with Atlantic distributions (Topham & Walker, loc. cit.). In the 

Azores it occurs in a more or less identical community on Cryptomeria. 

Apart from the type locality in Japan, it is known only from western Britain, the Azores, and 

the Canary Islands (Tenerife). It should be sought for in suitable habitats in other oceanic parts 

of Europe (e.g. south-west Norway, Bretagne, the Pyrenees, and Portugal).


Map 24 Micarea globulosella ® 1950 onwards + Micarea synotheoides # 1950 onwards O Before 1950 

43. Micarea ternaria (Nyl.) Vezda 

(Figs 32B, 52; Map 25) 

Sched. Lich. Sel. Exsicc. 858 (1970). - Lecidea sabuletorum f. ternaria Nyl. in Not. Sallsk Fauna Fl. fenn. 

Forhandl. 8: 151 (1866). - Lecidea ternaria (Nyl.) Nyl. in Flora, Jena 60: 232 (1877). Type: USSR, 

Lapponia murmanica, Kola Peninsula, 'Olenji', 1861, N. I. Fellman (H-NYL 18682 -holotype! [t.l.c: 

no substances]). 

Thallus effuse, muscicolous, composed of scattered to confluent, convex to irregularly 

subglobose areolae [? or saxicolous and obsolete -see 'observations']. -Areolae cream-white to 

ash-grey, c. 80-300 /xm diam; in section with hyahne amorphous covering layer c. 7-10 /xm tall, 

and outermost hyphae with greenish (K— , HNO3-I- red) walls. Phycobiont micareoid, cells c. 

4—7)Ltm diam. 

Apothecia numerous, black, matt or slightly glossy, 0-15-1 mm diam; at first (below 0-3 mm 

diam) ± globose or turbinate, later expanding to become broadly convex and ± adnate, 

sometimes faintly marginate. Hymenium c. 60-70 /xm tall, merging imperceptibly into the 

hypothecium; upper c. 10-15 ixm (epithecium) dark aeruginose-green, K-, HNO3+ purplered; 

remaining (lower) part ± hyaline with dilute green vertical streaks. Asci clavate. Spores 

fusiform, sometimes slightly curved, (0-)l-3-septate, 13-22(-24)x 3-5-5 /am. Paraphyses 

numerous, sparingly branched below but richly branched in epithecium, often anastomosing, c. 

1-1-8 /Am wide; apices often slightly incrassate and surrounded by dense pigment, and up to 3 

fxm wide. Hypothecium up to 380 /am tall; upper third to half, dilute olivaceous or dilute dull


brown; lower part ± hyaline; hyphae hyaline c. 0-7-1 -5 fim wide, interwoven or becoming 

vertically or outwardly orientated towards the hymenium and excipulum; hyphae in upper part 

of hypothecium intermixed with inflated, short-celled ascogenous hyphae. Excipulum usually 

distinct, hyaline, or tinged aeruginose or olive-green in parts; hyphae radiating, branched and 

anastomosing, 0-7-1 -7 /xm wide. 

Pycnidia rather numerous, sessile, black, 100-140(-200) /xm diam, ostioles often gaping; 

walls dark olive-green (K— , HNO3-I- red) above and at the sides, but becoming hyaline towards 

the base. Conidia (mesoconidia) cylindrical or oblong-ellipsoid, sometimes faintly biguttulate 

and slightly constricted in the middle, 4-6-6-3x1-2-1-7 /itm. Macro- or microconidial states not 

found. 

Chemistry: Thallus K-, C— , PD— ; sections of apothecia C-; t.l.c: no substances. 

Note: The above description is based largely on Alaskan specimens (Thomson 9188). 

Observations: M. ternaria is very close to M. lignaria but differs in having shorter spores that 

are never more than 3-septate, and in completely lacking lichen substances. In addition its 

apothecia have a more flattened appearance (sometimes with a faint marginal rim), and a ± 

clearly defined excipulum when viewed in vertical section. It is superficially similar to dark forms 

of M. peliocarpa but that species contains gyrophoric acid (thallus and apothecia in section C-lorange-red), 

has a hyaline hypothecium, and is not known to have pycnidia which contain 

mesoconidia. 

Having found most herbarium specimens named 'ternaria' to belong to M. lignaria or M. 

peliocarpa I suspected the name to be a synonym of one of these two species. The holotype of 

Map 25 Micarea subleprosula (§) 1950 onwards -I- Micarea cf. ternaria # 1950 onwards O Before 1950


Lecidea milliaria f. ternaria is very fragmentary and my first impression was that it was a 

diminutive specimen of M. lignaria, even though I failed to obtain a positive reaction with PD. 

This opinion was changed by my subsequent examination of the Alaskan material which is 

identical to the holotype in all respects but larger and in better condition. 

Several problematical collections from the British Isles (Shetland Islands and western 

Ireland) are provisionally referred to M. ternaria. They were all collected in coastal districts on 

hard siliceous rocks and have a ± obsolete thallus (PD— in microscopical preparations), rather 

flattened apothecia, and 0-3-septate spores. The specimen from Fair Isle has pycnidia, 

containing mesoconidia5-2-6-8xl-2-l-8/x,m (Fig. 52C). The discovery of an arctic species, such 

as M. ternaria, in coastal Britain is not without precedent. Examples are Lecanora straminea 

found in the Shetland Islands, Flannan Islands, and St Kilda, and Bacidia subfuscula found on 

North Rona, the Fame Islands, and as far south as the north coast of Norfolk and the Scilly Isles. 

A better understanding of the variability in habit and habitat of M. ternaria in the arctic should 

help to clarify the taxonomic relationships between the arctic and British populations. In 

addition, relevant material should be sought in coastal Norway. 

Habitat and distribution: On mosses and plant debris on the ground in coastal regions of arctic 

Europe and North America (northern Alaska). The Alaskan material {Thomson 9188) was 

growing with Micarea turfosa and Siphula ceratites. Southern populations on rocks may be 

represented in the British Isles (see 'observations' above). 

44. Micarea tuberculata (Sommerf.) R. Anderson 

(Figs32A,34,51B;Map26) 

in Bryologist 11: 46 (1974). - Lecidea tuberculata Sommerf., Suppl. Fl. Lapp.: 160 (1826). Type: 

Norway, Nordland, Saltdalen, Fiskevaagmollen, iii 1822, S. C. Sommerfelt (O - lectotype!; UPS - 


Lecidea latens Taylor in Mackay, Fl. Hib. 2: 259 (1836). Type: Ireland, Wicklow, The Dargle, T. Taylor 

(BM- lectotype!). 

Lecidea botryocarpa Nyl. in Flora, Jena 48: 603 (1865). Type: USSR, Karelskaya ASSR, Karelia 

onegensis, 'ad Onegam, Kapselka, 1863, T. Simming (H-NYL 10766 -holotype!). 

Lecidea subinfidula Nyl. in Flora, Jena 52: 295 (1869). Type: Finland, Lapponia enontekiensis, Naimakka, 

29 viii 1867, J. P. Norrlin 656 (H - lectotype!; UPS - isolectotype!). 

Lecidea tuberculata var. scandinavica Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 309 (1934). Type: Finland, 

Tavastia australis, Lammi, Evo, Lapinkallio, 1866, Norrlin (H-NYL 10767 - lectotype!). 

Thallus effuse, minutely scurfy-granular, or sometimes forming an irregularly rimose crust up 

to 0-2 mm thick, more rarely forming irregularly verrucose areolae c. 0-06-0-2(-0-3) mm diam; 

greenish buff, pale buff or greenish white; thin wefts of white prothalline hyphae sometimes 

visible. Phycobiont not micareoid; cells ± globose, thin-walled, 5-10(-12) /xm diam, or 

irregularly ellipsoid and up to 15x7 /xm. 

Apothecia numerous, convex-hemispherical and immarginate from the start, often becoming 

± globose or tuberculate, black and often with bluish tinge, matt, 0-16-0-3(-0-4) mm diam, or 

0-24-0-55 mm diam when tuberculate. Hymenium 30-35 jxm tall, dilute green or aeruginose, 

K- or + intensifying, HNO3+ purple-red, often with darker vertical streaks due to the presence 

of stout, pigmented paraphyses. Asci cylindrical-clavate, 25-30x7-9 /xm. Spores oblong-ovoid 

or oblong-ellipsoid, 0-1-septate, 5-5-8(-9)xl-5-2-5 /xm. Fflrap/iy:?^^ rather scanty, of two types 

(Fig. 34): p.p. evenly distributed, irregularly flexuose, simple or sparingly branched, often 

anastomosing, thin, 1-T5 /xm wide, sometimes widening to 1-7 /xm above, walls hyaline 

throughout and without adhering pigment; /?./?. fewer in number, as scattered individuals or in 

small fascicles, straight, simple or occasionally forked above, stout, coated ± throughout by 

dark greenish pigment and appearing c. 2-3 /u-m wide, apices sometimes ± incrassate and up to 4 

/xm wide (including pigment). Hypothecium c. 80-115 /xm tall, aeruginose- or olive-black, K-laeruginose 

intensifying; hyphae coated with dense dark green pigment and 2-3 /xm wide 

(overall), interwoven but becoming vertically orientated towards the hymenium and some


continuing into it as stout, pigmented paraphyses; ascogenous hyphae similarly pigmented, with 

swollen cells c. 3-5 fxm wide. Excipulum not evident, even in sections of young apothecia. 

Pycnidia often present, ± immersed, black, 60-120 /u,m diam, ostioles often gaping; walls dark 

aeruginose, K— , HNO3+ red; conidiogenous cells slender, cylindrical, 5-10x1-1-5 /u,m, often 

with a swollen, sometimes pigmented, base up to 3-2 jxm wide, and sometimes with one or two 

percurrent proliferations; conidia (mesoconidia) ± cylindrical, sometimes faintly biguttulate, 

3-4-3 X 1-1 -4 /xm. 

Chemistry: Thallus K— , C— , KC— , PD-; sections of apothecia C—; t.l.c: no substances. 

Observations: Micarea tuberculata is characterized by its rather small, markedly convex, often 

tuberculate, blackish apothecia, blue-green hymenium, aeruginose-black hypothecium and 

small, ovoid-oblong, 0-1-septate spores. The relative proportion of simple to 1-septate spores 

can vary greatly between collections, and in some specimens no septate spores can be found. M. 

tuberculata has often been confused with forms of M. sylvicola with small, immature spores, but 

such spores are always ellipsoid or ovoid-ellipsoid, and quite distinct from the predominantly 

oblong-ovoid spores of M. tuberculata. If pycnidia are present the two species can be separated 

on the size of their conidia, which are significantly longer (c. 4-6)u,m) in M. sylvicola. M. olivacea 

(q.v.) is superficially similar to M. tuberculata, but has slightly larger spores, shorter conidiogenous 

cells, and a micareoid phycobiont. Also of similar appearance is Psilolechia clavulifera 

(p. 201), but it has smaller spores, a pale hypothecium, numerous, stout, non-pigmented 

paraphyses, and a very different phycobiont. 

Map 26 Micarea tuberculata • 1950 onwards O Before 1950


Habitat and distribution: M. tuberculata is found on rocks, stones, and tree roots, etc. in dry, 

sheltered underhangs, and is a faithful member of the Micareetum sylvicolae. In the British Isles 

it has scattered localities in the north and west; although not yet known from the mainland in 

England and Wales it is expected to occur in those areas in suitable terrain (e.g. the Lake 

District, Dartmoor and Snowdonia). It is undoubtedly much overlooked, but appears to be 

genuinely rarer than M. bauschiana and M. sylvicola, with which it often occurs. 

It appears to be widely distributed in Fennoscandia where it is known from as far north as 

Nordland in Norway and Lycksele Lappmark in Sweden. It seems to be rare elsewhere in 

Europe, but I have seen material from southern Germany, northern Italy, and the Tatra 

mountains of Czechoslovakia. 

Exsiccata: Arnold Lich. Exs. 1057 (BM ex K, M). Larb. Lich. Herb. 221 p.p. (BM). Rabenh. Lich. Eur. 

648 p.p. (BM). Rasanen Lich. Fenn. 512 (BM, BM ex K), 672 p.p. (BM, LD-mixed with M. lutulata, 

M-mixed with M. sylvicola). 

\ 

45. Micarea turfosa (Massal.) Du Rietz 

(Fig. 33; Map 27) 

in Svensk hot. Tidskr. 17: 94 (1923). - Biatora turfosa Massal., Ric. Lich. Crost.: 128 (1852). Type: 

specimen unlocalized but probably from Sudety (Sudeten Mountains) in SW Poland, 'Flotow Lich. exs. 

130! hb. Flotow' (VER-holotype!). 

Biatora turfosa* verrucula Norman in K. nor. Vidensk. Selsk. Skr. 5: 353 (1868). - Lecidea verrucula 

(Norman) Th. Fr., Lich. Scand. 2: 523 (1874). - Micarea verrucula (Norman) Hedl. in Bih. K. svenska 

VetenskAkad. Handl. Ill, 18 (3): 84, 95 (1892). Type: Norway, Nordland, 'supra Heminghyt convallis 

Bejeren',/. M. Norman (-lectotype!; isolectotypes: LD!, M!, O!). 

Thallus effuse but often forming rounded patches about 3-5 cm diam, thin, ± uneven but not 

forming distinct areolae or goniocyst-like granules, blackish grey or brown-black but shaded 

portions sometimes dull grey-green, often appearing ± gelatinous when moist. Thallus in 

section up to 70 /xm thick, ecorticate and without an amorphous covering layer; outermost 

hyphae 2-4 /am wide with walls thickened by dark green pigment, K-, HNO3+ red; internal 

hyphae hyaline, c. 1-1-5 /xm wide. Phycobiont micareoid, cells 4-7 jxm diam. 

Apothecia numerous and often confluent, immarginate, convex to ± globose, black, matt or 

slightly glossy, rarely brown (shade forms), 0-15-0-3(-0-4) mm diam. Hymenium 35-50 îm tall; 

upper part (epithecium) sordid green, K— , HNOs-l- red, or sometimes brownish in part; middle 

part dilute aeruginose or olivaceous; lower part dilute reddish brown and merging ± impercep- 

tibly into the hypothecium. Asci clavate, 35-50 x 11-12 /xm. Spores oblong-ellipsoid to fusiform, 

sometimes slightly curved, simple or 1-3-septate, (10-)12-21(-25)x(3-5-)4-5 /itm. Paraphyses 

numerous, branched and sometimes anastomosing, c. 1-1-5 /xm wide in mid-hymenium but 

upper 5-15(-25) fxm thickened with dark greenish pigment and then l-5-2-5(-3) jxm wide. 

Hypothecium c. 70-140 /xm tall, mottled reddish brown, K-, HNO3- or turning orange-brown 

(never with purple tinge); hyphae interwoven but becoming vertically orientated towards the 

hymenium, hyaline or sometimes loosely coated with brown pigment, c. 1-5-1-7 /xm wide; 

ascogenous hyphae with swollen cells up to 5 ^tm wide. Excipulum reflexed but distinct, reddish 

brown but darker than the hypothecium; hyphae radiating, branched and anastomosing, 

l-l-5(-2) /xm diam, hyaline (pigment confined to gel-matrix). 

Pycnidia often present but very inconspicuous, immersed, 35-40 /u.m diam, wall dark sordid 

green, K— , HNO3+ 

red. Conidia (microconidia) ± cylindrical, c. 3-5-4-7x1 /xm. 

Chemistry: Sections of thallus and apothecia K- , C- 

, PD- 

; no substances detected by t. I.e. 

Observations: M. turfosa is fairly constant in appearance, except that the thallus is less well 

developed when in boggy habitats. Microscopically there is much variation between collections 

with regard to spore septation: some having mostly simple spores (e.g. Vezda Lich. Sel. 538), 

others having numerous 1-septate and several 2- or 3-septate spores (e.g. Vezda Lich. Sel. 

1135). In the type of Biatora turfosa the spores are mostly simple but a few with a single septum 

were found. In the type collections of Biatora turfosa* verrucula most spores are 1-septate.


With the combination of blackish thallus and black, convex to ± globose apothecia, M. 

turfosa is easily confused in the field with peat inhabiting forms of M. melaena, but the latter 

differs microscopically in having dark purple and/or green pigments in the hypothecium and 

thinner paraphyses. M. botryoides has similar apothecial pigmentation to M. turfosa, but its 

hypothecium is darker with nearly all the hyphae densely coated with pigment; in addition, M. 

botryoides has less numerous and (when hyaline) thinner paraphyses, a shallower hymenium, 

smaller spores, and its apothecia are always accompanied by numerous, stalked, black pycnidia. 

M. melaenida is unlikely to be mistaken for M. turfosa because of its whitish thallus and habitat 

of fine-grained, mineral soils; it can be further distinguished by its purple-brown epithecium, 

excipulum, and (usually) upper hypothecium. The little known M. osloensis differs from M. 

turfosa in several features but is most easily distinguished by its much smaller, non-septate 

spores. 

Habitat and distribution: M. turfosa seems to have an arctic-alpine distribution. In Britain it is 

known from summits in the Cairngorm, Grampian, and Breadalbane mountains of Scotland, 

where it grows on exposed turf (mostly over dead bryophytes) at altitudes of about 860-1245 m. 

In the Sudeten and Carpathian mountains of Czechoslovakia it is reported at altitudes of 1400 m 

and 1870 m respectively. It seems to be able to occur at lower altitudes in Sphagnum-bogs in 

Scandinavia, but no altitude data is given with any of the specimens seen. I have not seen any 

material from the Alps but it is likely to occur there. 

On the Cairngorm plateau M. turfosa occurs amongst Juncus trifidus. Associated lichens on 

the British gatherings include Cladonia spp., Lecidea caesioatra, and Lepraria neglecta. 

Map 27 Micarea turfosa • 1950 onwards O Before 1950


Outside Europe M. turfosa is little known, but I have seen material from Greenland and 

northern Alaska. 

Exsiccata: Flotow Lich. 130 (VER). Korber Lich. Sel. 12 (L). Magnusson Lich. Sel. Scand. 282 (GZU). 

Malme Lich. Suec. 865 (WIS). Norrlin & Nyl. Herb. Lich. Fenn. 321 (BM, M). Vezda Lich. Sel. 538 (BM), 

1135 (BM). 

Excluded taxa 

Bacidia beckhausii Korber, Parerga lich.: 134 (1860). - Micarea beckhausii (Korber) Vezda in Poelt & 

Vezda, Bestimmungsschl. europ. Flechten, Ergdnzungsheft I: 162 (1977). Type: Germany, 'West- 

phalen', Beckham (L 910, 137 1363 - lectotype!). 

Bacidia miniuscula Anzi, Cat. lich. Sondr. : 70 (I860). - Micarea miniuscula (Anzi) Vezda, in Vezda & V. 

Wirth in Folia geobot. phytotax., Praha 11: 100 (1976). Type: not seen. 

Bacidia beckhausii must be excluded from Micarea, mainly on account of its excipulum structure (see 

p. 189 and 198). 

Bacidia nitscltkeana var. perpusilloides Erichsen in Annals mycol. 41: 206 (1943). Type: West Germany, 

Schleswig-Holstein: Flensburg, Forst Clusries, near Wasserleben, on Picea twigs, 6x1923, C. F. E. 

Erichsen (HBG - lectotype!). Paratypes: Schleswig-Holstein: Flensburg, Jerrishoer Holz, 10 vi 1928, 

C. F. E. Erichsen (HBG!); Hamburg, Wohldorfer Wald, 12 vi 1905, C. F. E. Erichsen (HBG!); 

Hamburg, Volksdorfer Wald, 19 xii 1909, C. F. E. Erichsen (HBG!). All on young twigs of Picea. 

Thallus thin, green-grey. Apothecia minute, ± pellucid, pale brown, immarginate, c. 0-1-0-15 mm diam. 

Hymenium, hyaline, c. 30-40 /Am tall. Hypothecium dark brown, K-, or -I- greenish in part, c. 40-45 /im 

tall. Excipulum thin, c. 12-15 /xm wide, ± pseudoparenchymatous with cells c. 3-5-8x2-5-3 /Am (in K), 

hyahne or with greenish pigment in zone adjacent to hymenium. Asci clavate, numerous, 8-spored. Spores 

oblong-fusiform, straight or slightly curved, 3(-5)-septate, 16-24(-28)x3-4 /^m (Fig. 56A). Pycnidia often 

numerous, blackish, c. 50 /Am diam, wall greenish in K; conidia curved or sigmoid 12-30x0-8 /tm. 

Phycobiont cells c. 5-10 /im diam. 

This variety agrees in all respects, except host substratum, with Bacidia myrtillicola Erichsen in Mitt. 

Inst. allg. Bot. Hamb. 10: 414 (1939); type: Schleswig-Holstein: Lauenburg, 'im "Sachsenwald" bei 

Friedrichsruh an Vaccinium myrtillus am Rande des Reviers "Saupark".', 2 xi 1924, C. F. E. Erichsen 

(HBG - holotype!). B. myrtillicola belongs to the mainly foliicolous groups of Bacidia s. lat. that includes 

B. fuscatula (Miill. Arg.) Zahlbr., B. rhapidophylli (Rehm) Zahlbr., and B. vezdae Coppins & P. James. 

Catillaria melanobola f. frullaniae B. de Lesd., Rech. Lich. Dunkerque, Suppl. 1: 119 (1914). Type: 

France, Nord, near Burgues, on Frullania on Ulmus, M. B. de Lesdain, Zahlbr. Lich. Rar. Exs. 152 (BM 

-isotype!). 

This is an apparently undescribed (at specific rank) species of Arthonia s. lat. The apothecia are very 

small, c. 0- 1 mm diam, convex-globose and black. In section the epithecium and hypothecium are brownish 

(K-l- olive), the asci clavate, c. 21 x 10 /tm at maturity, with a minute internal amyloid ring [as described for 

Bryostigma leucodontis (Poelt & Dobbeler, 1979) and Chrysothrix (Laundon, 1981), and noted by me in 

several Arthonia species, especially lichenicolous species and members of the subgenus Allarthonia], 

spores hyaline, 1-septate, 9-11x2-3-3 /im, and paraphyses scanty, thin, c. 0-6-1 /Am wide. This species is 

rather common in the British Isles where it is usually referred to as 'Arthonia cf. exilis\ It is found in a 

variety of habitats, for example bark of shaded trunks of Acer, Fraxinus and Ulmus, branches and twigs of 

Corylus and Sambucus, over bryophytes (especially Frullania) on trunks of Quercus and Ulmus, tufts of 

Armeria on sea-cliffs, and possibly also on sandstone rocks. When on bark the apothecia may attain a larger 

size, to 0-2 mm diam. Corticolous material has recently been distributed as Vezda Lich. Sel. 1701. In E it is 

represented by material from the following British vice-counties: 1, 20, 29, 31 , 62, 64, 67, 78, 83, 88, 90, 98, 

99, 110, H18 and H20; and from Denmark and Germany (Schleswig-Holstein). 

There has been much confusion regarding the correct interpretation of Arthonia exilis (Florke) Anzi. I 

have recently had the opportunity of examining type material of this species (Florke Deutsch. Lich. 187 

(WRSL - lectotype!)) and a brief description of it is as follows: 

Thallus lignicolous (probably on worked timber; accompanied by a few granular-areolae of Candelariella 

vitellina), thin, grey-white. Phycobiont noX Trentepohlia, cells, 10-19 /Am diam. Apothecia numerous, ± 

regularly dispersed, black, 0-1-0-2 mm diam. Epithecium (including lower border with substratum) 

reddish brown, K-. Hymenium c. 30 /im tall, I -I- reddish, K/I-l- blue. ^5d clavate, with minute amyloid 

ring ("Bryostigma -type'), 22-30x14-15 /im, 8-spored. Spores hyahne, 1-septate, 8-12x2-5-3-5(^) /Am. 

Paraphyses ± coherent (even in K), rather stout, c. 1-5-2 /xm wide; apices ± swollen with brown


Fig. 56 A, spores of Bacidia myrtillicola (HBG - lectotype of Bacidia nitschkeana var. perpusilloides). 

B-C, Catillaria bouteillei (H-NYL 1884 - holotype of Lecidea littorella); B, spores; C, conidiogenous 

cells and conidia. Scale = 10 /u,m. 

pigment-caps; closely adhering pigment sometimes continuing down to about mid-hymenium level. 

Hypothecium hyaline. 

The status oiA. exilis s. str. as a British species requires confirmation. 

Catillaria rhodosphaeraTh . Fr. & HultinginTh. Fr. Lich. Scand. 2: 571 (1874). Type: Sweden, Dalsland, 

Haverud, 1870, /. Hulling (UPS - lectotype! ; UPS 

- isolectotype!). 

This species was provisionally referred to Micarea by Kilias (1980: 392), but in fact belongs to Biatora 

Fr., a name which requires conservation against Biatora Ach. (= Stenhammarella Hertel). In my opinion 

C. rhodosphaera represents a saxicolous form of Catillaria sphaeroides (Dickson) Schuler. 

Lecidea arceutina var. hypnaea Nyl. in Flora, Jena 51: 165 (1968). - Bacidia arceutina f . hypnaea (Nyl.) A. 

L. Sm. , Monogr. Brit. Lich. 2: 158 (1911). Type: Jersey, 1866, Larbalestier {W-]


arceutina (Ach.) Arnold. In his diagnosis Nylander gives the spore length as 45-70 /xm and on the holotype 

packet he wrote '45-20x1 yam'; my examination of the holotypefound the spores to be 38-51x1-5 /xm. 

Lecidea clavulifera Nyl. - see Psilolechia clavulifera. 

Lecidea demarginata Nyl. in Flora, Jena 61: 245 (1878). Type: Finland, Tavastia australis, Evo, 'supra 

saxum in sylva juniori, loco olim deusto', 1873,7. P. Norrlin, Norrlin & Nyl. Herb. Lich. Fenn. 179 (Hlectotype!; 

isolectotypes (without exsiccate label): H-NYL 20162! & 20163!). 

In the protologue Nylander cited material of Norrlin's from Finland and of Larbalestier's from Ireland. 

The epithet 'demarginata' was first used in 1875 on the label of Herb. Lich. Fenn. 179, although without a 

diagnosis. An example of this exsiccate is here selected as lectotype, and is a specimen of Lecidea erratica 

Korber. The cited Larbalestier material is represented in BM and H and belongs to M. lutulata. 

Lecidea denigrata var. bacidiella Vainio in Medd. Soc. Fauna Fl. fenn. 10: 28 (1883). - Micarea bacidiella 

(Vainio) Vezda, in Vezda &. V. Wirth in Folia geobot. phytotax., Praha 11: 100 (1976). Type: Finland, 

Lapponia kemensis, Sodankyla, Pyhatunturi, Kannolla, on hgnum, 1878, E. A. Vainio (TUR-VAINIO 

22505 -holotype!). 

As stated by Vainio (1934: 462) this is a synonym of Bacidia miniuscula Anzi (i.e. B. beckhausii Korber). 

The following were notes made from the holotype of var. bacidiella: 

Thallus lignicolous, entirely endoxylic. Phycobiont cells c. 8-14 ^tm diam. Apothecia thinly marginate 

and ± plane when young, soon becoming convex and immarginate, black, c. 0-2-0-3 mm diam. Hymenium 

c. 35 /xm tall, with olivaceous vertical streaks, K+ violet, C+ violet (not red). Asci clavate, 30-35x10-12 

/im, 8-spored. Spores rod-shaped or slightly curved, (l-)3(-7)-septate, 17-26x1-7-2 /nm. Paraphyses 

simple or sparingly branched, 1-1-5 /xm wide; apices somewhat incrassate, to 2 /xm wide. Hypothecium 

hyaline. Excipulum dilute olivaceous (K+ violet), of radiating, branched hyphae c. 1-5-2 /xm wide 

embedded in a dense gel; hyphae becoming more distinct in K but not separating. Pycnidia immersed in the 

substratum, black, c. 50-100 /xm diam; walls olivaceous, K-l- violet. Conidia simple, hyaline, oblong- 

ellipsoid, often wider at proximal end, 2-8-3-5 x 1(-1 -4) /xm. 

B. beckhausii usually occurs on bark, although I have seen a few additional lignicolous specimens from 

Scotland. Micarea bacidiella sensu Vezda & Wirth {loc. cit. ) is not B. beckhausii but the superficially similar 

Micarea globulosella (see p. 135). 

Lecidea erysiboides Nyl. in Not. Sdllsk. Fauna Fl. fenn. 4: 232 (1859). - Catillaria erysiboides (Nyl.) Th. Fr. 

Lich. Scand. 2: 572 (1874). Type: Finland, Nylandia, Helsingfors [Helsinki], 'Gumtackt' on rotting 

decorticate pine trunk, 1858, W. Nylander (H - lectotype!; H-NYL p.m. 4514- isolectotype!). 

This name has been misapplied by many lichenologists and the majority of specimens seen belong to 

Micarea prasina, or other species such as Catillaria sphaeroides, Lecania cyrtella, and Micarea denigrata. 

The type material of L. erysiboides is not a Micarea but a species of Catillaria s. lat. , although I am 

uncertain of its affinities. It has small, plane to convex, reddish brown (testaceous) apothecia that are 

marginate when young, the excipulum is composed of much-branched, radiating hyphae which are distinct 

in K but still tightly bound by the gel matrix, and the spores are ovoid and often constricted at the septum, 

1-septate with the upper cell usually enlarged and ± globose, 8-9-5x3-5 /xm. C. erysiboides has not been 

correctly reported from the British Isles but it should be looked for, especially in the eastern Scottish 

highlands. 

Additional specimens of C. erysiboides examined: NORWAY. Hordland: Granvin, on Betula lignum, 

1904, Havaas, Lich. Exs. Norv. 292 (BG). FINLAND. Nylandia: Helsinki, 1859, W. Nylander (H-NYL 

21650), and 1861 (BM). ITALY: Trentino ('Sudtirol'): Paneveggio, on top of cut stump of Picea, 2 ix 1883, 

Arnold, Lich. Exs. 1002 (BM ex K). USSR. 'Lapponia orientalis', 1863, A^. /. Fellman, Lich. Arct. 156 

(BMexK). 

Lecidea fuliginea Ach., Syn. Lich.: 35 (1814). - Micarea fuliginea (Ach.) Fr., Syst. orb.: 257 (1825). - 

Micarea fuliginea (Ach.) Fr., Stirp. agrifemsion. : 37 (1825); comb, inval. (Arts 34.1, 43). 

I have not seen the type of this name but it is probably a synonym of Lecidea icmalea Ach. A later 

synonym of the latter is Pannularia perfurfurea Nyl. which is the type of the genus Placynthiella Gyelnik. 

Lecidea gelatinosa Florke in Magazin Ges. naturf. Fr. Berl. 3: 201 (1809). - Micarea gelatinosa (Florke) 

Brodo in Bryologist 70: 216 (1967). 

I have not seen type material of this species, but if its general interpretation is correct then it is not a 

Micarea. In fact it belongs in the Lecidea granulosa group, which includes L. aeruginosa Borrer, L. 

aeneofusca Florke ex Flotow, L. granulosa (Hoffm.) Ach. and L. viridescens (Schrader) Ach. These 

species are not congeneric with the type species of Lecidea Ach. (L. fuscoatra (L.) Ach.) and they await the 

formal transference to a genus in the Trapeliaceae . Further detailed studies are needed to see if they can be 1


placed in Trapelia Choisy or Trapeliopsis Hertel & G. Schneider, or if a new genus is required to 

accommodate them. 

Lecidea littorella Nyl. in Flora, Jena 60: 229 (1877). - Catillaria littorella (Nyl.) Zahlbr., Cat. lich. univ. 4: 

56 (1926). Type: Ireland, West Galway, 'Hibernia occidentalis, Bord du Lough Inagh, mais extremement 

rare', 1876, C. Larbalestier (Yi-^Yl. 18884- holotype!, isotypes: BM!, BM ex K!). 

The following notes were made from the holotype: Thallus saxicolous, not delimited but forming small 

patches amongst Hymenelia lacustris and Porina chlorotica, whitish, or dull ochraceous in part (? due to 

age), rimose, matt. Apothecia numerous, weakly marginate, plane to slightly convex, pallid to dull 

orange-red, 0-1-0-3 mm diam; margin not exceeding the level of the disc, very thin, c. 0-1-0-3 mm diam; 

margin not exceeding the level of the disc, very thin, c. 0-01-0-02 mm wide, whitish with faintly pruinose 

appearance. Hymenium 40-47 jxxn tall, hyaline. Asci cylindrical-clavate, ' Lecanora-type\ 8-spored. 

Spores ovoid to ovoid-oblong, often 'slipper-shaped', 1-septate and often constricted at the septum, 

9-5-14(-16)x4-6(-7) ^tm (Fig. 56B). Paraphyses numerous, thin, 0-8-1 ixm wide, simple or sparingly 

branched below, becoming wider (to 1-7 ixm) and more frequently branched above. Hypothecium hyaline. 

Excipulum hyaline or dilute yellow-straw, inspersed with minute crystals in water mounts but ± clearing in 

K, c. 20 /am wide laterally, slightly widening below to 28 )u,m, minutely paraplechtenchymatous with cells in 

the lower part measuring 4—6x2-5^ /u,m. Pycnidia frequent, at first immersed, later becoming emergent, 

white (translucent when wet), c. 80-100 fxm diam. Conidia (Fig. 56C) simple, ovoid to pyriform, 

2-8xl-5-l-7)Ltm. 

L. littorella was provisionally referred to Micarea by Kilias (1981: 392). However, my subsequent 

examination of the holotype proved it to be a saxicolous form of the normally foliicolous Catillaria 

bouteillei (Desm.) Zahlbr.; previous reports of this species on rock are given by Degelius (1944) and 

Santesson (1952). C. bouteillei is apparently widely distributed in Ireland, especially on the leaves oiBuxus 

(Knowles, 1929; Scannell, 1978). An additional British saxicolous specimen has been located: V.C.5, 

South Somerset, Broomfield, Ruborough Camp, 1914, W. Watson (BM ex K). 

Nylander's choice of the epithet 'littorella' is rather misleading because the holotype was not collected on 

the sea-shore, but on the shores of an inland, freshwater lake some 10 km from the sea. British authors (e.g. 

James, 1970; Fletcher, 1975) have mistakenly applied the name Catillaria littorella to a species that grows in 

crevices in siliceous rocks near the sea-shore; this species probably belongs to the perplexing Lecania 

ery5/fte complex. 

Lecidea milliaria a. [var.] terrestrisFr. Lich. Eur. 342 (1831). Type: not designated. 

This name is of unlikely appHcation in M/carea. Fries cited his L/c/i. Suec. no. 213, the example of which 

in UPS is a member of the Lecidea limosa group. 

Lecidea ocelliformis Nyl. in Flora, Jena 48: 145 (1865). - Bilimbia ocelliformis (Nyl.) Branth. & Rostr. in 

Bot. Tidskr. 3: 226 (1869). - Lecidea atroviridis f. ocelliformis (Nyl.) Hedl. in Bih. K. svenska 

VetenskAkad Handl. Ill, 18 (3): 64 (1892). - Catillaria prasina f. ocelliformis (Nyl.) Erichsen in Annls. 

mycol. 41: 205 (1943). Type: Finland, Tavastia australis, Hollola, ad corticem Sorbi, 30 vi 1863, /. P. 

Norrlin 210 (H-NYL 20607! - labelled Tsotype' by M. Inoue in 1979). 

This is not a Micarea. If the types of L. atroviridis (Arnold) Th. Fr. and L. ocelliformis are conspecific, as 

placed by Hedlund (op. cit.) and Vainio (1934: 218), then the latter epithet has priority. 

Lecidea pauxilla Krempelh. in Verh. zool.-bot. Ges. Wien 26: 455 (1876). Type: New Zealand, on bark 

[? Podocarpus], 18 -, C. Knight (M - lectotype!; M - isolectotype!). 

L. pauxilla was given as a synonym of Catillaria synothea (i.e. Micarea denigrata) by Zahlbruckner (Cat. 

lich. univ. 4: 78, 1926). However, the type material belongs to Cliostomum griffithii (Sm.) Coppins. 

Lecidea recondita Erichsen in Annls mycol. 42: 25 (1944). Type: Germany, Schleswig-Holstein: 'Apen- 

rade, an Blocken am Grunde einer Erosionsschluct im Geholz Jiirgensgaard' [now Denmark, Jylland, 

near Abenra], 28 vii 1932, C. F. E. Erichsen (HBG - holotype!). 

In the protologue Erichsen compares this with Lecidea sylvicola (= Micarea sylvicold). My examinations 

of the holotype proved it not to be a Micarea but referable to Catillaria chalybeia (Borrer) Massal.; see 

Kilias (1980: 448). 

Lecidea sabuletorum f. simp/icior Nyl., Lich. Scand.: 205 (1861). - Bilimbia trisepta f. simplicior (Nyl.) 

Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 258 (1922). Type: not designated. 

Nylander (loc. cit.) cited the following material 'supra muscos ad Helsingforse [ipse] et in Sueciae 

montibus [Thedenius] atque supra terram in Lapponia'. 

According to Fries (1874: 523) the 'Helsingforse' specimen is Lecidea verrucula (= Micarea turfosa); 

according to Vainio (1922: 140) the Swedish specimen from Funnesdalsberget in Harjedalen collected by


K. F. Thedenius is Lecidea dufourei (= Catillaria contristans (Nyl.) Zahlbr.). I am uncertain as to the 

identity of the specimen(s) referred to by Nylander as 'in Lapponia'. This may be based on the specimen 

cited by Vainio (1922: 140, 258) as 'supra Grimmias in Kipina Lapponiae a G. SeHn coUecta', which Vainio 

refers to Bilimbia trisepta [sensu Vainio = Micarea peliocarpa] f. simplicior. However, material in 

Nylander's herbarium (H-NYL 18841) labelled 'Lapponia, G. Selin 1861' is Catillaria contristans. 

Furthermore, another specimen of 'Lecidea sabuletorum f. simplicior' collected from Iceland by Isaac 

Carroll in 1861 (H-NYL 18862) is also C. contristans. It would appear that Nylander's concept of 

'simplicior' is best fitted by C. contristans and should be typified on the material collected by Thedenius or 

Selin. 

Lichen niger Hudson, Fl. angl., ed. 2, 2: 524(1778). -Collemanigrum{Hudson)Y{offm. , Deutschl. Fl.: 103 

(1796). - Micarea nigra (Hudson) Fr., Syst. orb.: 257 (1825). - Micarea nigra (Hudson) Fr., Stirp. agri 

femsion.: 37 (1825); comb, inval. (Arts 34.1, 43). - Placynthium nigrum (Hudson) Gray, Nat. Arr. Br. 

PI. 1:395(1821). 

This is the type species of the genus Placynthium Gray. 

Lichen viridescens Schrader, in Gmelin Syst. nat. 2 (2): 1361 (1792). - Lecidea viridescens (Schrader) Ach. 

Meth. Lich. : 62 (1803). - Micarea viridescens (Schrader) Brodo in Bryologist 70: 216 (1967). 

I have not seen the type of this name but if its normal interpretation is correct then the species belongs to 

the Lecidea granulosa group; see notes above under Lecidea gelatinosa. 

Micarea chrysophthalmaF . James in Lichenologist 5: 131 (1971). - Chrysothrix chrysophthalma (P. James) 

P. James & Laundon, in Laundon in Lichenologist 13: 104 (1981). 

This is a species of Chrysothrix and is discussed in detail by Laundon (1981). 

Micarea coccinea Fr., Syst. orb. : 

257 (1825); nom. nudum (Art. 32.1). 

I do not know the identity of this species, and I have not seen this name in any other publication. On 

morphological grounds it is unlikely that Fries intended a new combination based on Lichen coccineus 

Dickson [= Haematomma ochroleucum (Necker) Laundon] or Lecidea coccinea Schw. [= Lecidea russula 

Ach.]. 

Micarea cyanescens Poelt & Dobbeler in Bot. Jb 96: 339 (1975). Holotype: Germany, Bayern, Altbayerische 

Alpen, Chiemgauer Berge, 900-1000 m on Campylium halleri on north-facing calcareous rocks, 29 

ix 1974, /. Poelt (GZU, not seen). Paratypes: Austria, Tirol, Lechtaler Alpen, 'unteres Medriol-Tal 

uber Zams bei Landeck, an Dolomitblocken unter Griinerlen', 1600-1700 m, on Campylium halleri, 

1969, J. Poelt (hb Poelt 13295!); Austria, Steiermark, 'uber Kalk, Rote Wand bei Mixnitz, c. 1470 m, 

unweit des Gipfels', 1974, J. Poelt (GZU!). 

This is a very curious species; its hymenium components are embedded in a dense gel which does not 

disperse in 50% of KOH. It is not a Micarea and a new genus is probably required to accommodate it. 

Micarea hylocomii Poelt & Dobbler in Bot. Jb. 96: 341 (1975). Type: Austria, Tirol, Samnaun Gruppe, on 

the way from Serfaus to Madatschen, c. 1500 m, on Hylocomium splendens, 15 ix 1972, J. Poelt (GZU - 

holotype!). 

I endorse the describing authors' view that the position of this species in Micarea is doubtful, and I 

beheve a new genus should be erected to accommodate it. 

Micarea leprosa P. James in Lichenologist 5: 133 (1971). - Vezdaea leprosa (P. James) Vezda, in Poelt & 

Dobbeler in Lichenologist 9: 170 (1977). 

This is a species of Vezdaea (Poelt & Dobbeler, 1975). A large collection of it from Scotland has recently 

been distributed as Hertel Lecid. Exs. 60. 

Micarea minima Poelt & Dobbeler in Bot. Jb. 96: 342 (1975). Holotype: Germany, Rheinland-Pfalz, 

'Boschung eines Waldweges bei Battweiler, Kreis Zweibriicken, c. 400 m', on Polytrichum commune 

and P. formosum, 1974, J. Poelt (GZU, not seen). Paratype: Austria, Karnten, Nockgruppe, Afritzer 

Berge, 'Bergwald und saure Weiden iiber Verditz, nordlich Villach', c. 1300 m, on Polytrichum 

formosum, 1974,/. Poe/f (hb Poelt 13294!). 

I was unable to find any apothecia in the examined paratype and so I cannot confidently exclude this 

species from Micarea. Further studies are required to establish its generic disposition. 

Patellaria nigrataMuW. Arg. in Flora, Jena7l: 540(18SS).-Bacidianigrata{Mu\\. Arg.) Zahlbr., Car. lich. 

univ. 4: 129 (1926). - Micarea nigrata (Miill. Arg.) Kalb, Lichenes Neotropici Fasc. 1: 8 (1981); comb, 

inval. (Art. 33.2). Type: Brazil, Apiahy, on argillaceous soil, 1883, Puiggari (G!). 

This species closely resembles a Micarea and even appears to have 'micareoid' algae. However, its


excipular hyphae are conglutinated, broad (c. 2-2-5 /u,m in K), and pachydermatous. It is closely allied to 

Bacidia subrudecta (Vainio) Zahlbr. and may have affinities with Byssoloma, but I think it is best retained 

in Bacidia s. lat. pending further study. The material distributed by Kalb {Lich. Neotrop. Exs. ^2) as 

'Micarea nigrata' is M. lignaria var. lignaria. 

Psilolechia clavulifera (Nyl.) Coppins, comb. nov. - Lecidea clavulifera Nyl. in Flora, Jena 52: 294 (1869). - 

Micarea clavulifera (Nyl.) Coppins & P. James, in D. Hawksw. , P. James & Coppins in Lichenologist 12: 

107 (1980). Type: Finland, Lapponia kemensis, Muonionska [Muonio], Keimioniemi, ad radicem 

abietis, 1867, J. P. Norrlin 627 (H - lectotype!; isolectotypes: H!, H-NYL 20855!). 

Lecidea clavulifera f. subviridicans Nyl. in Flora, Jena 60: 463 (1877). Type: Ireland, West Galway, 

Connemara, Kylemore, in a cave on the NW side of Doughraugh mountain, 1876, C. Larbalestier (BMlectotype! 

Isolectotypes: BM ex K!; also distributed as Larbal. Lich. Herb. 29: BM!, H!). 

Thallus effuse, whitish or greenish white, of dispersed to coalescing, irregular granular-areolae c. 

Fig. 57 Psilolechia clavulifera (Coppins 3614, E). A, spores. B, paraphyses. C, phycobiont cells (cell 

contents and mycobiont hyphae omitted). Scale = 10 /xm.


0-1-0-2 mm diam. Areolae often disintegrating to form a scurfy-granular crust. Phycobiont not micareoid; 

cells in clusters and tightly bound by short-celled hyphae, but haustoria not observed; clusters interconnected 

by filamentous hyphae c. 1-5-2 ju.m wide. Phycobiont cells irregularly globose, broadly ellipsoid or 

oblong, c. 5-12(-18)x3-8 />tm, often arranged in pairs or in short chains of up to four cells (Figs. 57C). 

Apothecia convex-hemispherical and immarginate from the beginning, sometimes becoming subglo- 

bose, more rarely tuberculate, grey-black with bluish tinge, but shade forms sometimes whitish, blue-grey 

or grey-brown, 0-l-0-3(-0-4) mm diam; base of apothecia with a white rim (c. 50 /xm wide) of outwardly 

radiating hyphae. Hymenium 28-35 /xm tall, dilute straw (shade forms), or dilute greenish or aeruginose 

(K-l- green intensifying) especially in the upper part. Asci cylindrical-clavate, 25-30x5-7 /im, 8-spored. 

Spores ovoid, oblong-ovoid or dacryoid, simple, 3-6x(l-)l-2-l-7(-2) /xm (Fig. 57A). Paraphyses (Fig. 

57B) numerous, usually branched, sometimes anastomosing, distinctly septate and appearing ± articu- 

lated, stout, 1-3-2 /Ltm wide; apices sometimes ± clavate and up to 3 jxm wide, walls not pigmented but 

often surrounded by deeply pigmented gel matrix. Hypothecium c. 50-70 /tim tall, hyaline, or dilute 

greenish or aeruginose (K— , HNO3-I- red) but then never darker than the hymenium. Excipulum 

indistinct, of radiating hyphae that protrude as loose, hyaline hyphae c. 1-5-2 /xm wide and up to 50 /x,m 

long. 


Chemistry: All parts K- , KC- , C- , PD- ; no substances detected by t.l.c. 

Preliminary studies of Lecidea clavulifera by Mr P. W. James and myself led us to transfer it to Micarea. 

However, critical studies have given me second thoughts on this placement. The distinct white rim of 

protruding excipular hyphae (superficially like those of Byssoloma spp.), stout and distinctly septate 

paraphyses, and the unusual phycobiont are all uncharacteristic of a Micarea. The consideration of 

Psilolechia Massal. as an alternative genus for L. clavulifera was prompted by observations of a 

lichenicolous member of the CaHciales, Microcalicium arenarium (Hampe ex Massal.) Tibell; M. arenarium 

is usually found as a parasite of Psilolechia lucida (Ach.) M. Choisy (see Tibell, 1978), but at three 

localities in Scotland I have found it on L. clavulifera. At the Berwickshire locality the host and parasite 

occurred in abundance on roots, stones, and soil of two up-ended trees (Fraxinus); P. lucida was also 

present in quantity but was not parasitized by M. arenarium. 

With the exception of pigmentation, sections of the ascocarps of P. lucida and L. clavulifera show ± 

identical anatomical features, e.g. nature of paraphyses, size and shape of asci and spores, and excipulum 

with numerous protruding hyphae. P. lucida differs from P. clavulifera in the yellow-green colour of its 

leprose-granular thallus and apothecia (due to presence of pulvinic acid derivatives), and a different 

phycobiont with ± globose cells 5-14 /xm diam. The thallus hyphae of both species are identical in 

appearance, and pycnidia are not known in either species. A recent account of P. lucida is given by James 

(in Poelt & Vezda, 1981). The phycobiont of P. clavulifera is very unusual; it is reminiscent of the 

Stichococcus phycobiont of species such as Chaenotheca stemonea and Coniocybe furfuracea, but its cells 

are much larger. 

P. clavulifera occurs in communities of the Micareetum sylvicolae, on roots, stones, and consolidated soil 

of underhangs on banks or the root systems of up-ended trees. Most British records are from Scotland, but 

it is also known from Cumbria, south Wales and western Ireland (Map 28). From outside Britain I have 

seen material of it from Finland, Germany, and Czechoslovakia (see below). 

Additional specimens of P. clavulifera examined: British Isles. Brecon (V.C.42). 22/94: Upper 

Dyfnant Valley, 1982, Woods (hb Woods). Cumberland (V.C.70). 35/54: Baron Wood, by River Eden, 

60-75 m dry sandstone cliff, 1979, Coppins 4344 (E). Berwick (V.C.81). 36/76: W of Elba, S side of 

Whiteadder Water, exposed roots of up-ended Fraxinus, 1981, Coppins 8890 (E - Microcalicium 

arenarium folder). West Perth (V.C.87). 27/40: Aberfoyle, the Trossachs, on stone amongst roots of 

up-ended tree, 1978, Coppins3661 (E). Mid Perth (V.C. 88). 27/55: Black Wood of Rannoch, stone in bank 

by track, 1976, Coppins 4654 (E); 27/81: Crieff, Drummond Wood, roots and sandstone of up-ended tree, 

partly parasited by Microcalicium arenarium, 1978, Coppins 3164, 3625 (E). Angus (V.C. 90). 37/33: 

Sidlaw Hills, Auchterhouse Hill, 396 m, underside of boulder, 1975, Coppins 846 (E - Microcalicium 

arenarium folder). South Ebudes (V.C. 102). 16/49: Colonsay, Coille Mhor, roots and soil of up-ended 

Betula, 1981, Coppins 8878 (E). Mid Ebudes (V.C. 103). Mull: 17/32: Bunessan, Ardfenaig Woods, 1970, 

James (BM); 17/54: Salen, 1968, James (BM); 17/55; Aros House, roots of up-ended tree, 1968, James 

(BM). West Galway (V.C.H.16). 84/72 (L/65): Connemara, near Clifden, 1878, Larbalestier (BM). 

Finland. Satakunta: Siikainen, Vuorijarvi, Vaasaneva, sandy soil amongst roots of Picea, 1936, Laurila 

(GZU, H). W. Germany. Baden-Wiirttemberg: Heidelberg, Konigstuhle, on sandstone, 1883, Zwackh 

(H-NYL p.m. 4229). Czechoslovakia. Slovakia, Nizke Tatry, Liptovska Teplicka, Dzurova, 18 -, Lojka 

4289 (BM).


Map 28 Psilolechia clavulifera # 1950 onwards O Before 1950 

Sporacestra Massal. in Atti R. 1st. veneto Sci. Ill, 5: 264 (1860). Type species: Biatora prasina Tuck. & 

Mont, in Mont. (1857), non (Fr.) Trevisan (1856). 

Sporacestra was included as a synonym of Micarea by Vezda & Wirth (1976: 99) who confused the two 

quite separate applications of the name 'Biatora prasina' . The name Biatora prasina Tuck. & Mont, was 

introduced for a corticolous species from Venezuela with long acicular spores, and no reference was made 

to the 'prasina' of Fries. Similarly, Massalongo made no reference to Fries' name and his generic 

description (e.g. '. 

. . sporidii capillari aghiformi, lineari allungati . . .') clearly refers to the stated type 

species, ' Biatora prasina Mont., Tuck.'. 

The Venezuelan species was transferred to Bacidia de Not. by Zahlbruckner, but his use of the epithet 

'prasina' is contrary to the present Code. From the descriptions of this species I believe it should be 

retained in Bacidia, pending further study. The nomenclature and required new combination for the 

species is as follows: 

Bacidia prasinata (Tuck.) Coppins, comb. nov. - Biatora prasinata Tuck, in Syn. N. Am. Lich 2: 41 (1888); 

nom. nov. - Bacidia prasina Tuck & Mont, in Mont, in Annls Sci. nat. IV, 8: 296 (1857), non (Fr.) Trevisan 

(1856). - Bacidia prasina Zahlbr., Cat. lich. univ. 4: 253 (1926); nom. illeg. (Art. 11). Type: Venezuela, on 

bark, Fendler (not seen). 

Index to exsiccatae 

I have attempted to examine at least one example of all European and North American material 

referable to Micarea and distributed in recognized exsiccatae (Lynge 1915-22, 1939; Sayre, 

1969). My preliminary list was compiled from Lynge {op. cit.), available schedae, major floras


(e.g. Fries, 1874; Korber, 1855, 1859-65; Smith, 1926; Vainio, 1922, 1934) and numerous other 

papers dealing with species here included in Micarea. Most examples gleaned from these sources 

have been seen; the few not seen are listed after the main index. Herbarium locations for 

examined examples containing Micarea species can be found in the relevant taxonomic 

accounts; locations of examples not containing Micarea species are given in the index. The 

names on the exsiccate labels are given (in round brackets following the number) where they are 

taxonomically at variance with the species represented. 

In many cases, only one or two examples of a particular number have been seen by me, thus 

this index must be regarded as provisional. Some numbers listed as not containing a Micarea 

(e.g. Zwackh Lich. Exs. US) may well in further examples (not seen by me) contain a Micarea. 

In addition, some Micarea species may have been misidentified and distributed under names not 

connected with Micarea, and consequently overlooked in the course of this study. 

It should be noted that many numbers in the exsiccatae of Harmand {Lich. Loth.), Mougeot 

& Nestler, and Schaerer are notoriously heterogeneous with regard to species compo- 

sition, substratum, and locality. Most numbers in other exsiccatae are reasonably uniform in 

these respects but heterogenicity sometimes arises in cases where species of similar outward 

appearance occur together in the field (e.g. M. bauschiana-M. lutulata-M. sylvicola-M. 

tuberculata, and M. denigrata-M. lignaria-M. peliocarpa). During the preparation of exsiccatae 

the need for the careful identification of individual samples is paramount before they are dis- 

tributed. 

Generic names are abbreviated as follows: Bacidia (B), Biatora (Bi.), Biatorina (Biat.), 

Bilimbia (Bit.), Catillaria (C), Lecanora (Lee), Lecidea (L.), Micarea (M.), Psilolechia {P.) 

and Scoliciosporum (5.). 

ANZI, Lich. Exs. Ital.: 256 M. denigrata; 259A (Bil. lignaria) M. melaena; 259B {Bil. hypnophila - given 

by Lynge as Bil. lignaria) B. sabuletorum (BM). 

ANZI, Lich. Lang.: 148 {Bil. syncomistd) M. lignaria, one example in BM with a little M. peliocarpa. 

ANZI, Lich. Sondr.: 170A M. peliocarpa; 170B M. melaena. 

ARNOLD, Lich. exs.: 120 M. bauschiana; 167A {Bil. lignaria ^saxigena Leighton) M. peliocarpa; 167B 

M. peliocarpa; 217 M. nitschkeana; 279, 280A-C M. prasina; 332A-C M. melaena; 348A, B M. lignaria; 

409A M. sylvicola; 409B (L. sylvicola) M. sylvicola, or M. lutulata; 503A-D M. nitschkeana; 548, 549 M. 

cinerea; 556[A], B (L. assimilata) M. crassipes; 626, 627 M. misella; 836 M. lithinella; 837 M. peliocarpa; 

1051 {Bil. ternaria) M. peliocarpa; 1057 M. tuberculata; 1121 M. crassipes; 1122 M. prasina; \A1\ M. 

elachista; 1472 M. prasina. 

ARNOLD, Lich. Mon.: 46 M. denigrata; 47 M. cinerea; 48 M. nitschkeana; 49 M. melaena; 115, 116 M. 

cinerea; 118 {Bil. trisepta) M. peliocarpa; 172 M. misella; 241 M. misella; 243 M. prasina; 244 {Biat. 

prasiniza) M. adnata; 245 M. prasina; 246 M. elachista; 248, 249 M. melaena; 269 {Bil. trisepta) M. 

peliocarpa; 270 {Bil. trisepta) M. nitschkeana; 307 M. misella; 357 {Bil. trisepta) M. peliocarpa; 407 M. 

melaena; 482 {Bil. trisepta) M. peliocarpa. 

BOHLER, Lich. Brit.: 85 (L. viridescens) M. lignaria. 

BRITZELMAYR, Lich. exs.: 174 {Biat. prasiniza var. laeta) M. adnata; 175 {Biat. glomerella) L. 

viridescens (M); 208 M. misella; 310 {Biat. synothea) L. turgidula + Lee. sp. (H); 464 {Biat. synothea) M. 

denigrata + C. nigroclavata (H); 599 {Biat. synothea) Arthonia vinosa (H); 829 {Bil. milliaria f. nigrita) 

M. nitschkeana; 846 M. cinerea; 946 M. melaena; 961 (L. assimilata f. irrubata) L. hypnorum + L. limosa 

(H). 

CLAUDEL & HARMAND, Lich. Gall. : 43 M. lignaria; 89 M. nitschkeana; 445 M. misella. 

CROMBIE, Lich. Brit.: 174 (L. misella) M. misella +/or M. prasina. 

CUMMINGS, Decad. N Amer. Lich. Ed. I: 302 M. lignaria; 355 M. prasina. Ed. II: 232 M. lignaria. 

ELENKIN, Lich. Ross.: 189 M. misella. 

Erb. Critt. Ital., ser. I: 198 (C. synothea) C. nigroclavata (UPS). 

FELLMAN, Lich. Arct.: 159 M. melaena; 164 (L. assimilata f. alpestris) M. incrassata, + a little M. 

crassipes in example in H; 165 (L. assimilata) M. crassipes; 166 (L. limosa) M. incrassata. 

FLAGEY, Lich. Franche-Comte: 137 {Bi. synothea) Lee. sp. + Lecidella ? euphorea (UPS). 

Flora Hung, exs.: 714 M. melaenida. 

FLOTOW, Lich. exs.: 129A M. lignaria; 129B (L. milliaria b. ma/o/- - according to Lynge) Mycoblastus 

affinis (UPS); 129C (L. milliaria var. lignaria - according to Lynge) M. melaena; 129D (L. milliaria) C. 

globulosa (UPS); 129E M. lignaria; 130 M. turfosa; 131 M. lignaria; 171A M. sylvicola.


FRIES, Lich. Suec: 29 M. lignaria, + some M. denigrata in example in UPS; 98 M. denigrata, + some M. 

lignaria in example in E; 212A, B M. melaena. 

HANSEN, Lich. Groenl.: 172 (L. assimilata) L. stenotera (BM); 247 (L. assimilata) L. limosa (BM). 

HARMAND, Guide Elem. Lich.: 91 M. lignaria. 

HARMAND, Lich. Loth.: 810 (Lecidea vernalis, and f. prasina) mixture from several localities, example 

in ANGUC has Bi. (? Moelleropsis) humida, C. sphaeroides, L. icmalea and L. vernalis, that in UPS has 

C. sphaeroides and L. vernalis; 838 M. denigrata; 852 M. lignaria; 853 M. nitschkeana. 

HAVAAS, Lich. Norv.: 139 M. subviolascens; 555 M. lignaria; 571 (L. sylvicola var.) M. lutulata; 694 M. 

subviolascens; 710 M. subviolascens. 

HAVAAS, Lich. Norv. Occid.: 269 M. subviolascens. 

HEPP, Flecht. Eur.: 14 M. denigrata; 20 {Bi. lignaria) M. nitschkeana; 21 (Bi. cinerea) M. cinerea, or M. 

nitschkeana; 278 M. prasina; 284 M. peliocarpa; 285 (Bi. lignaria var. milliaria) M. peliocarpa; 504 M. 

melaena; 510 M. peliocarpa; 524 {Bi. asserculorum) S. umbrinum (E). 

HEPP, Zurich: 206 {L. cinerea) M. peliocarpa; 210 (L. synothea) Lecania cyrtella agg. (BERN); 224 M. 

prasina. 

HERTEL, Lecid. exs.: 34 M. lignaria; 54 M. ? bauschiana. 

JOHNSON, N. Engl. Lich. Herb.: 373 M. denigrata; 375 (L. turneri; listed under Bil. lignaria by Smith 

(1911, 1926)) Bacidia sabuletorum + Toninia lobulata (BM); 376 M. melaena; 434 L. sylvicola var. 

hellbomii) M. sylvicola, or M. bauschiana; 453 M. lignaria; 504 M. bauschiana. 

KALB, Lich. Neotropici: 22 M. lignaria (M. nigrata); 186 M. lignaria. 

KAVINA & HILITZER, Crypt. Cech.: 269 M. sylvicola. 

KORBER, Lich. Sel. Germ.: 12 M. turfosa; 75 M. sylvicola; 133A {Bil. lignaria) M. peliocarpa; 133B M. 

peliocarpa; 137 {Bi. denigrata) L. (Lecidella) alba (L); 250 M. prasina. 

Krypt. Exs. Vind.: 165 M. peliocarpa; 362 M. melaena; 658 M. lignaria; 1232 M. nitschkeana; 1532 M. 

misella; 2061 (C prasiniza var. prasinoleuca) Dimerella diluta (BM, BM ex K, M); 2268 M. assimilata; 

2561 (C denigrata) C. globulosa (BM, BM ex K); 3153 M. denigrata; 3154 M. melaenida; 3651 (L. 

misella) M. denigrata; 4214 M. misella; 4858 M. denigrata. 

KUTAK, Lich. Bohem. : 205 (L. asserculorum) M. denigrata; 310 M. prasina; All M. lignaria, + a little M. 

leprosula in example in O; 516 M. denigrata; 517 M. denigrata. 

LARBALESTIER, Lich. Caesar. Sarg.: 83 (L. arceutina var. hypnaea) M. lignaria; 84 M. sylvicola. 

LARBALESTIER, Lich. Herb.: 29 P. clavulifera; 68 M. bauschiana; 223 M. lutulata; 227 (L. polioides) 

M. tuberculata, but example in LIV is L. monticola;212 M. lignaria; 304 M. sylvicola; 305 M. sylvicola or 

M. bauschiana; 347 M. peliocarpa. 

LEIGHTON, Lich. Brit.: 120 (Bi. uliginosa) M. melaena; 210 M. lignaria; 238 (L. milliaria var. terrestris) 

M. lignaria, or M. peliocarpa, example in MANCH has a little M. leprosula; 386 M. lignaria; 388 (L. 

sabuletorum var. milliaria) M. botryoides. 

LOJKA, Lich. Hung.: 60 M. cinerea; 61 M. lignaria; 134 M. peliocarpa. 

LOJKA, Lichenoth. Univ.: 29-31 M. prasina; 137 M. nitschkeana; 233 M. lithinella. 

MAGNUSSON, Lich. Sel. Scand.: 134 M. prasina; 208 M. leprosula; 282 M. turfosa; 340 M. nitschkeana. 

MALBRANCHE, Lich. Norm.: 287 (L. sphaeroides var. lignaria) M. nitschkeana; 387 (L. sphaeroides 

var. melaena) M. denigrata. 

MALME, Lich. Suec: 20 M. rhabdogena; 21 M. elachista; 22 M. anterior; 23, 24 M. prasina, 25 M. 

nitschkeana; 26 M. eximia; 27 M. melaena; 28 M. contexta; 125 M. lithinella; 145 M. denigrata; 169 M. 

peliocarpa; 199 M. sylvicola; 216 M. assimilata; 288 M. lignaria; 362 M. crassipes; 365 (M. denigrata var. 

pyrenothizans) M. misella; 865 M. turfosa. 

MIGULA, Crypt. Germ., Aust., Helv.: 1 (5/7. milliaria) M. lignaria +lox M. leprosula, ± M. nitschkeana; 

132 M. denigrata; 226 M. lignaria. 

MOUGEOT & NESTLER, Stirpes Crypt. Vog.-Rehn.: 1329 (L. melaena) an example in E includes a little 

A/, melaena, other collections in BM (2 sets) , DBN and M (3 sets) contain a selection from the following, 

Lee. spp., L. granulosa agg., L. turgidula, L. sp., M. denigrata; 1430 M. lignaria. 

MUDD, Lich. Brit.: 156-8 M. lignaria; 159 M. melaena; 164 (L. prasina) L. viridescens (E, BM, M); 175 

M. sylvicola. 

NORRLIN & NYLANDER, Herb. Lich. Fenn.: 145 (L. sylvicola) L. conferanda (BM, H, M); 177 M. 

denigrata; 180 M. melaena; 182 (L. pelidna) M. intrusa [some examples may prove to be S. umbrinum]; 

194A, B M. crassipes; 314 M. elachista, one example H is B. hegetschweileri; 319A, B (L. milliaria var. 

ternaria) M. lignaria; 321 M. turfosa; 724 M. elachista; lAA M. misella; 745 (L. asserculorum) M. 

denigrata; 763, 764 (L. tuberculata) M. sylvicola. 

OLIVIER, Lich. Orne: 237 M. melaena; 344 A/, lignaria. 

PISUT, Lich. Slov.: 156 M. //gnana.


POELT, Lich. Alp.: 22 (L. misella) M. denigrata. 

RABENHORST, Lich. Eur.: 224 {Bi. turfosa) L. uliginosa (BM ex K); 322 M. lignaria, example in M has 

some M. peliocarpa; 582 {Bil. lignaria) M. nitschkeana; 583 M. nitschkeana; 603 {Bil. syncomista) M. 

lignaria; 626 M. denigrata; 648 {Bi. bauschiana) M. bauschiana, or M. tuberculata; 675 (L. sylvicola) L. 

erratica (BM, M); 676 M. prasina; 733 M. prasina. 

RASANEN, Lich. Fenn. exs.: 489 {Bil. nitschkeana) B. naegelii (LD); 512 M. tuberculata; 642 M. 

nitschkeana; 651, 652, 653 M. prasina; 672 (L. sylvicola) M. sylvicola, +/or M. tuberculata, +/or M. 

lutulata; 822 (C elachista f. simplicata) L. turgidula (BM ex K); 963 M. melaena. 

RASANEN, Lichenoth. Fenn.: 137 (C. elachista f. simplicata) B. igniarii (BM ex K); 343 M. peliocarpa; 

344 M. melaena; 426 (C ///cw) M. globulosella. 

Reliq. Suza.: 42 M. misella. 

ROUMEGUERE, Lich. Gall.: 87 (L. vernalis var. milliaria) B. sabuletorum (BM, M); 193 (L. vernalis 

var. synothea) M. lignaria; 231 M. melaena, but example in BM is mixture of a L. sp. and Lee. spp.; 232 

M. lignaria; 467 (L. melonida fmelaenidaj) Toninia aromatica (BM). 

SAMPAIO, Lich. Port.: 132 M. denigrata; 147 M. lignaria. 

SCHAERER, Lich. Helv.: 196 (L. sabuletorum var. lignaria) examples labelled 'Ad ligna decorticata in 

alpibus' are M. prasina, examples labelled 'In m. Belpberg' contain one, or a mixture, of the following, 

M. lignaria, M. melaena, M. peliocarpa and M. leprosula; 327 (Parmelia varia var. denigrata) L. 

(Lecidella) alba (BM ex K), or L. aff. cadubriae (E). 

SUZA, Lich. Bohem.: 131 M. sylvicola. 

THOMSON, Lich. Arct.: 12 (B. melaena) Toninia lobulata (LD). 

VAINIO, Lich. Bras.: 1420 M. misella; 1451 M. misella. 

VEZDA, Lich. Bohem.: 133 M. lignaria; 258 M. lignaria; 282 M. crassipes. 

VEZDA, Lich. Sel. exs.: 11 M. crassipes; 14 M. melaena; 90 M. prasina; 516 M. lignaria; 538 M. turfosa; 

706 (M. violacea) B. naegelii (BM); 858 (M. ternaria) M. lignaria; 957 (L. tuberculata) M. sylvicola; 1036 

M. lignaria; 1087 M. cinerea; 1088(M. ternaria)M. lignaria; 1134M. elachista; 1135 M. turfosa; 1341 (A/. 

tuberculata) M. sylvicola; 1342 M. peliocarpa; 1380 M. peliocarpa; 1430 M. denigrata; 1467 M. prasina; 

1595 (M. hemipoliella) M. prasina; 537 M. melaenida. 

WEBER, Lich. Exs.: 73 M. denigrata. 

ZAHLBRUCKNER, Lich. Rar. exs.: 110 M. nitschkeana; 152 (C. melanobola i.frullaniae =) Arthonia 

aff. exilis (BM); 174 (L. infidula) L. erratica; 175 M. prasina; 276 M. turfosa. 

ZWACKH, Lich. exs.: 121 M. lignaria; 122 (L. milliaria Fr. var. ?) M. elachista; 276 M. peliocarpa; 219 

(Bi. gelatinosa var. minorl) M. bauschiana; 279B M. bauschiana; 394 M. denigrata; 416 M. prasina; All 

(Bil. nitschkeana) S. umbrinum + S. chlorococcum (UPS); 470 M. nitschkeana [two labels: 'Bilimbia 

nitschkeana, bei Miinster in Westfalen'; or 'Lecidea nitschkeana, bei Dolbrueck in Kreise Paderborn 

. . .', sometimes labelled '470 bis\ example in M. also has M. denigrata]; 534 M. nitschkeana; 535 (L. 

latens) M. sylvicola; 587 M. nitschkeana; 590 M. lithinella; 591 A, B, 592A-E, 593A-C M. prasina; 594A, 

B, 595 M. bauschiana; 596 (L. latens) M. sylvicola; 597 M. sylvicola; 656 M. prasina; 657 M. melaena; 

675 M. melaena; 118 (L. cinerea) L. symmicta agg. (UPS); 780 (L. trachona) M. sylvicola; 897 M. 

peliocarpa; 898 M. cinerea; 900 M. turfosa; 919 M. sylvicola; 1085 M. misella. 

Examples of the following have not been examined by me; names taken from Lynge (op. d/.). 

BRITZELMAYR, Lich. exs.: 838 {Bil. lignaria); 890 {Bit. trisepta). FLOTOW, Lich. exs.: 112 (L. synothea ?). 

HARMAND, Lich. Loth.: 834 (L. denigrata). KORBER, Lich. Sel. Germ.: 343 {Bit. syncomista). LARBALESTIER, 

Lich. Herb.: 139 (L. micrococca). OLIVIER, Lich. Orne: 264 (L. nitschkeana). ROUMEGUERE, Gen. Lich. exs.: 36 

(L. vernalis var. synothea); 51 (L. vernalis var. milliaria). No examples of this exsiccata have been traced; it is almost 

certain that it was never distributed (P. M. J0rgensen, in litt.). SCHADE, STOLLE & RIEHMER, Lich. Sax.: 256 {B. 

trisepta). ZWACKH, Lich. exs.: 131 (L. glomerella and L. confluens). 

Acknowledgements 

My sincerest thanks go to Mr P. W. James and Dr F. Rose for their encouragement, guidance and many 

kindnesses, not only during the present study, but also from the time of my first interests in hchenology in 

1965. For further help and advice in those earher years I here take the opportunity of extending my thanks 

to Dr D. L. Hawksworth, Mr J. R. Laundon, Dr M. R. D. Seaward, Dr T. D. V. Swinscow, and Mr A. E. 

Wade. 

I am much indebted to the curators and owners of the herbaria listed in 'Materials'; especially as their 

searches for relevant.specimens and processing of loans have often been long and arduous procedures. 

Furthermore, many of these curators and collectors have provided me with much supplementary 

information on localities, habitats, and historical background. Special thanks are due to Miss F. J. Walker 

A


of the British Museum (Natural History) for her efficient handUng of my many 'urgent' requests for 

herbarium and bibliographical information. 

Recognition for their helpful comments and suggestions on various nomenclatural and taxonomic 

problems pertaining to Micarea and related groups goes to Mr B. L. Burtt, Prof. Dr H. Hertel, Dr H. 

Kilias, Prof. P. M. J0rgensen, Prof. Dr J. Poelt, Prof. R. Santesson, Dr V. Wirth, and Dr A. Vezda. 

Much advice and assistance has come from many of my colleagues at the Royal Botanic Garden, 

Edinburgh; in particular, I would like to thank Prof. D. M. Henderson, the Regius Keeper, and Mrs 

F. M. Bennell, Mr A. P. Bennell, Mrs N. M. Gregory, Dr R. Watling, Mrs J. M. Woods, Mrs H. Hoy, Mrs 

D. A. Morrison, Mr M. V. Mathew, Mr D. G. Long, and the staff of the typing section. 

All the people and organizations to whom I am grateful for assistance with field excursions are too 

numerous to mention individually. However, special thanks go to Dr L. Tibell for arranging and leading 

me on a tour of mid- and north Sweden in 1977, and to the Nordic Lichen Society (especially Mr U. 

S0chting and Mr K. Ramkaer) for inviting me as guest visitor to their excursion in north Jylland in 1979. 

The following funds and institutions are thanked for financial support of field-work: Nature Conservancy 

Council, Natural Environment Research Council, Royal Botanic Garden, Edinburgh, and the World 

Wildlife Fund. 

Finally, I must thank my wife and family for tolerating the many sacrifices that they have had to endure, 

especially during the preparation of the final manuscript. 

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Index 

Accepted names are in roman and synonyms in italic. New names and principal references are in bold 

whilst an asterisk (*) denotes a figure or map. Some taxa mentioned incidentally (e.g. in lists of associated 

species) are not included. 

aberrans 185, 186 

abietina (Lecanactis) 67 

absistens (Bacidia) 88 

adnata 24, 29, 33*, 61, 64, 65, 66*, 67, 68*, 89, 

90, 93-95, 105, 108-110, 111*, 114, 204 

aeneofusca (Lecidea) 198 

aeolotera (Huilia) 26 

aeruginosa 151 

aeruginosa (Lecidea) 198 

affinis (Mycoblastus) 204 

aggerata 186 

alabastrites 22, 23, 26*, 29, 32*, 34*, 64, 65, 85, 

89-91, 93-95, 99, 103, 110-113*, 122, 172 

alba (Lecidea, Lecidella) 205,206 

albella 121,122 

albicans 170 

albidolivens 170 

alnicola 151 

andesitica 128 

aniptiza 127 

anterior 24, 29, 32, 33*, 61, 64, 65, 69*, 90, 91, 

95, 100, 105, 108-110, 112-114, 136, 205 

antrophila 149 

aphanoides 138, 140 

apochroeella (Lecidea) 132 

arceutina (Bacidia) 197, 198 

arenarium (Microcalicium) 202 

arnoldiana (Bacidia) 67 

aromatica (Toninia) 206 

Arthonia 31,98 

Arthoniaceae 31 

Arthothelium 31 

asserculorum (Lecidea) 158, 159 

assimilata 22, 25, 29, 32, 35*, 61, 62, 64, 67, 

87-90, 95, 100, 104, 114-116*, 154, 186, 187, 

205 

atroviridis (Lecidea) 199 

Bacidia 88, 97 

bacidiella (Lecidea, Micarea) 135, 198 

bauschiana 24, 25, 29, 30*, 31, 36*, 61, 64, 71*, 

87, 89, 93, 95, 100, 105, 117-119*, 148, 149, 187, 

194, 204-206 

(A. 

beckhausii (Bacidia, Micarea) 20, 30, 88, 102, 

135, 189, 196, 198 

berengeriana (Lecidea) 115 

Biatora 97,112,197 

Biatorina 97 

biforme (Anisomeridium) 66 

biseptata \1A 

Bispora 29 

botryocarpa 192 

botryoides 24, 29, 32, 36* , 61 , 62, 64, 65* , 67, 

69*, 84, 88-90, 95, 99, 106, 108, 114, 118-121*, 

149, 156, 159, 163, 195, 205 

botryoides (Byssus) 174 

bouteillei (Catillaria) 197*, 199 

brasiliana 158 

Bryophagae 96, 196 

Bryostigma 31 

Buellia 31,61 

byssacea 174, 175 

Byssoloma 98, 202 

cadubriae (Lecidea) 206 

caesioatra (Lecidea) 30, 116 

calamophila 142 

callicarpa 117 

Caloplaca 31 

carbonicola 132 

carneoglauca (Bacidia) 67 

Catillaria 61,88,97 

catillarioides 151 

Chaenothecopsis 64 

chalybeia (Catillaria) 97, 199 

chlorococcum (Scoliciosporum) 91, 97, 206 

chrysophthalma (Chrysothrix, Micarea) 200 

Chrysothrix 98, 196 

cinerea 22-26, 29, 37*, 61, 63-66, 70*, 85, 87, 

88-91, 93-95, 99, 104, 108, 109, 112, 121-123*, 

124, 129, 141, 144, 166, 172, 204-206 

clavulifera (Psilolechia, Lecidea, Micarea) 105, 

107, 163, 193, 198, 201*, 202, 203*, 205 

Cliostomum 97 

coccinea (Micarea) 96, 200 

Collemaceae 20, 24

conferanda (Lecidea) 205 

confusula 183 

conglomerata 169 

contexta 24, 29, 31, 32, 38*, 61, 63, 64, 71*, 85, 

87, 88, 90, 91, 95, 100, 107, 124, 125, 134, 164, 

205 

contristans (Catillaria) 98, 115, 200 

contrusa 138 

crassipes 20, 28*, 29, 38*, 62-64, 67, 72*, 87-89, 

95, 96, 100, 106, 115, 116, 125, 126, 152, 187, 

204-206 

cupreola 170 

curvata 29, 38*, 64, 85, 87-89, 92, 95, 105, 126, 

127, 185 

cyanescens (Micarea) 200 

cyrtella (Lecania) 198, 205 

cyrtellina (Lecania) 66 

dedivitatum 174 

decrustata 158 

delicatula 121, 122 

demarginata (Lecidea) 149, 198 

denigrata 22, 23*, 24, 26, 29, 30, 32, 39*, 61, 64, 

66, 67, 73*, 84, 85, 87-90, 92-96, 99-102, 108, 

109, 127-130*, 131, 133-136, 148, 166, 182, 198, 

204-206 

diluta 112 

diluta (Dimerella) 205 

dilutiuscula 117 

discrepans 174 

discretula 111 

dufourei (Lecidea) 200 

ecrustacea Lamy 127 

ecrustacea Nyl. ex Vainio 181 

efflorescens (Lecidea) 85 

elachista 22, 23*, 29, 40*, 64, 84, 88, 90, 91, 95, 

99, 103, 105, 129, 131-133, 181, 204-206 

elegantior (Huilia) 26 

endocyanea 151 

endoleuca 64, 84-86, 93-95, 104, 122, 146, 147* 

epiphaeotera 151 

erratica (Lecidea) 67, 187, 198, 206 

erysibe (Lecania) 199 

erysiboides (Catillaria, Lecidea) 20, 114, 198 

euphorea (Lecidella) 204 

exigua 170 

exilis (Arthonia) 196, 197, 206 

eximia 24, 29, 40*, 61-64, 71*, 87, 90, 91, 95, 

100, 107, 124, 134, 164, 168, 205 

flotowii 186 

fraterculans 170 

friesiana 128 

friesiana (Argopsis) 85 

frullaniae (Catillaria) 196, 197, 206 

fucatus (Mycoblastus) 88 

fuliginea (Lecidea, Micarea) 96, 198 

fusca , spododes var . 165 


fusca, synothea f. 128 

fuscatula (Bacidia) 196 

Fuscidea 22,97,185 

fuscoatra (Lecidea) 97, 198 

fuscopurpurea (Arthonia) 31 

gelatinosa (Lecidea, Micarea) 198 

geomaea 142 

geophana (Steinia) 98 

glebosula VIA 

globifera 158 

globularis 49*, 158 

globulosa (Catillaria) 66, 204, 205 

globulosella 22, 23, 41*, 64, 74*, 85, 88, 90, 95, 

96, 99, 101, 134, 135, 166, 180, 189, 190*, 198, 

206 

glomerella 131 

gomphillaceum 63, 96, 143, 144 

graniforme (Cliostomum) 67 

granulosa (Lecidea) 87, 98, 198, 200, 205 

granvina 149 

Graphidaceae 31 

griffithii (Cliostomum) 67, 159, 199 

gymnomitrii (Bryomyces) 143 

hardangeriana 185 

hedlundii 24, 31, 40*, 64, 75*, 85, 88, 90, 91, 95, 

100, 102, 108, 114, 135-137, 140, 175 

hegetschweileri (Bacidia) 205 

hellbomii 186 

Helocarpon 96, 126 

hemipoliella 39*, 127, 131 

hemipolioides 170 

Herteliana 97 

Huilia 26, 97 

humida (Biatora, ?Moelleropsis) 205 

hylocomii (Micarea) 200 

hymenea (Pertusaria) 88 

hypnaea (Lecidea) 197, 198 

hypnorum (Lecidea) 31, 88, 115-116, 204 

hypocyanea 186 

hypoleuca 122 

hypopta (Lecidea) 132 

icmalea (Lecidea) 87 , 198 , 205 

igniarii (Bacidia) 206 

ilyophora 151 

incincta 186 

incrassata 22, 23, 25, 28*, 29, 42*, 61, 64, 67, 

72*, 88, 89, 95, 96, 100, 104, 115, 116*, 137, 138, 

204 

infidula 117 

infuscata 115, 137 

intrusa 20, 23, 25, 29, 32, 43*, 61, 64, 85, 87-89, 

95, 105, 138*, 139, 140, 184*, 185, 205 

irrubata 114 

juistense (Anisomeridium) 66, 67

212 

laeta 174 

lapiseda 170 

latens 192 

laxula 148 

Lecanora 61,88 

Lecanoraceae 98 

Lecidea 88, 97 

Lecideaceae 19, 31 , 71 , 87, 97, 98 

Lecidella 22,31,97 

leprosa (Vezdaea, Micarea) 200 

leprosula 22, 44*, 64, 85-87, 89, 90, 95, 99, 103, 

107, 140-142*, 172, 183, 205 

leucococca 170 

leucodontis (Bryostigma) 31, 196 

levicula (Lecidea) 87, 100 

lignaria 20, 22, 23*, 24, 27*, 29, 31, 45*, 61, 63, 

64, 66, 76*, 84-90, 92-96, 99, 104, 122, 126, 

129, 141, 142-145*, 146, 166, 170-172, 191, 192, 

197,201,204-206 

limosa (Lecidea) 98, 115-116, 199, 204, 205 

lithinella 24, 28*, 32, 46*, 61, 64, 85, 89, 92, 93, 

95, 100, 105, 118, 147, 148, 150*, 204-206 

littorella (Catillaria, Lecidea) 197* , 199 

livescens 170 

livida B. de Lesd. 165 

livida Korber 170 

lobariella (Dactylospora) 88 

lobulata (Toninia) 145 , 205 , 206 

longior 128 

lucida (Psilolechia) 202 

lutulata 24, 25, 29, 46*, 64, 65, 75*, 88, 89, 95, 

100, 106, 118, 120, 148-150*, 161, 187, 198, 

204-206 

lynceola 117 

major 128 

malmeana 134 

meizospora 143 

melaena 22, 29, 47*, 61-64, 77*, 85, 87-96, 100, 

103, 107, 124, 150-153*, 154, 165, 171, 187, 195, 

204-206 

melaenida 22, 24, 29, 48*, 64, 88-90, 92, 95, 100, 

106, 115, 154, 155, 195, 204-206 

melaeniza 24, 29, 47*, 64, 65, 69*, 85, 90, 91, 95, 

99, 106, 108, 114, 120, 155, 156, 159, 161, 164 

melanobola 24, 31, 47, 61, 64, 77*, 85, 88, 90, 95, 

100, 102, 156, 157, 175 

melanochroza 158 

Melanolecia 97 

melaphana 43*, 138 

meridionalis 154 

Metacapnodiaceae 67 

Micarea 20, 96-100 

Micareaceae 98 

microcarpa 165 

micrococca 173-176 

microspora 117 

milliaria 84, 142 

minima (Micarea) 200 


miniuscula (Bacidia, Micarea) 196, 198 

misella 24, 29, 31, 49*, 61, 64, 65, 67, 85, 88, 89, 

90, 94-96, 100, 102, 107, 114, 120, 156, 157, 

158-160*, 164, 175, 181, 182, 204-206 

monticola (Lecidea) 205 

moriformis 125 

muhrii 24, 29, 49*, 61, 64, 88-91, 95, 106, 148, 

149, 156, 160, 161 

myriocarpa 24, 29, 31, 32, 49*, 62, 64, 78*, 

88-90, 95, 105, 106, 148, 149, 161, 162*, 163 

myrtillicola (Bacidia) 196, 197* 

naegelii (Bacidia) 172, 206 

niger (Lichen) 200 

nigra (Micarea) 96, 200 

nigra, melanobola f. 174 

nigra, spododes var. 165 

nigella 24, 29, 31 , 50* , 61-64, 78* , 87, 90, 91 , 94, 

95, 100, 107, 108, 114, 120, 124, 134, 156, 159, 

163, 164*, 165 

nigrata Miill. Arg. (Bacidia, Micarea, 

Patellaria) 200,201,205 

nigrata Nyl. 143 

nigroclavata (Catillaria) 204 

nigrum (Collema, Placynthium) 200 

nitschkeana 22, 23, 30, 50* , 64, 66, 67, 85, 

87-96, 99-101, 129, 135-136, 165-167*, 171, 

172, 189, 204-206 

niveoatra (Opegrapha) 66, 67 

Nostoc 25,96,114,155 

obscurior 117 

occulta \1A 

ocelliformis (Lecidea) 199 

oculata (Pertusaria) 88 

olivacea 24, 29, 31, 50*, 62-64, 78*, 84, 89, 90, 

95, 100, 107, 124, 134, 164*, 165, 167, 168, 193 

osloensis 29, 31, 54*, 61, 64, 88, 89, 95, 106, 169, 

195 

pallida (Lecidea) 26 

panaeola (Huilia) 25, 140 

parissima 127 

paucula 149 

pauxilla (Lecidea) 199 

peliocarpa 22-25, 27*, 29, 51*, 61, 63-66, 79*, 

85-87, 89-91, 93-96, 99, 103, 104, 108, 109, 

111, 112, 122, 129, 141, 144, 149, 166, 169-173*, 

191,200,204-206 

perfurfurea (Pannularia) 198 

perpusilloides 196, 197* 

phaeobaea (Arthonia) 67 

Phyllopsora 22, 85 

Placynthiella 198 

poliococca 131 

poliococcoides 131 

poliodes 148 

polytrichi 53*, 80*, 96, 174, 176 

polytropoides 22 , 45 * , 143

populina (Lecidea, Micarea) 31,32 

praeviridans 127 

prasina Fr. 20, 23*, 24-26, 29, 30, 32, 52*, 53*, 

61, 64, 66, 80*, 85-96, 100, 102, 108, 109, 129, 

136, 148, 157, 173-176*, 177*, 178*, 179, 198, 

204-206 

prasina Tuck. & Mont. (Bacidia, Biatora) 203 

prasinata (Bacidia, Biatora) 203 

prasiniza 53*, 109, 173 

prasinoleuca 174 

pruinosum (Scoliciosporum) 30, 97, 180 

pseudoglomerella 128 

Psilolechia 98, 202 

pulverula 125 

pycnidiophora 23, 29, 54*, 64, 65, 67, 85, 89, 90, 

93, 95, 99, 103, 108, 114, 179, 180*, 181, 182 

pyrenothizans 111 

quercicola (Lecanora) 66 

radiata (Arthonia) 31 

recondita (Lecidea) 199 

relicta 154 

rhabdogena 24, 29, 31, 55*, 64, 85, 88, 90, 91, 95, 

99,105,133,164,181,182,205 

rhapidophylli (Bacidia) 196 

rhodosphaera (Catillaria) 197 

Rhopalospora 97 

rosei (Phyllopsora) 85, 146 

rosella (Bacidia) 97 

rusticella 117, 118 

sabuletorum (Bacidia) 145, 204-206 

salevensis (Bacidia) 171 

sanguineoatra (Lecidea) 115 

saprophila 151 

saxkola 142 

saxigena Hepp 169, 170 

saxigena Leighton 143 

scandinavica 192 

schadeanum (Scoliciosporum) 180 

schumannii 48*, 154 

Scoliciosporum 97, 98, 103, 140 

semialbula 128 

semipallens 117 

simplicata 131 

simplicior (Lecidea, Bacidia) 199, 200 

smaragdina 186 

sordidescens 173, 174 

sororians 131, 133 

spadicea (Arthonia) 67 

sphaeroides (Bacidia, Bilimbia, Catillaria, 

Lichen) 20, 112, 197, 198, 205 

spodiza 111 

spododes 165 

Sporacestra 203 

Steinia 98 

Stenhammarella 197 

stenotera (Lecidea) 98, 115-116, 205 


Stereocauliscum 96 

Stigonema 25,96,114,115 

stipitata 23 , 29 , 55 * , 64 , 65 , 67 , 89 , 91 , 93-95 , 99 

103, 108, 114, 180*, 182 

stizenbergeri 151 

straminea (Lecanora) 192 

Strangospora 98 

subabietina (Lecanactis) 67 

subassimilata 125 

subdiscordans (Byssoloma) 98 

subfuscula (Bacidia) 192 

subinfidula 192 

subleprosula 20, 24, 31, 32, 56*, 64, 85-87, 89, 

94, 95, 99, 103, 107, 141, 182, 183, 191* 

sublivescens 111 

sublivida 186 

submisella 127 

subnigrata 22, 23, 29, 32, 57*, 61, 64, 67, 81*, 

88-90, 95, 105, 127, 139, 183, 184*, 185 

subrudecta (Bacidia) 201 

subviolascens 20, 22, 23, 25, 29, 57*, 64, 88, 89, 

95,100,101,115,185,186,205 

subviridescens 52*, 173, 174, 177 

subviridicans 201 

sylvicola 20, 23-25, 29, 31, 57*, 61, 62, 64, 66, 

82*, 87-90, 93, 95, 96, 100, 107, 118, 149, 152, 

186-188*, 193, 194, 199, 204-206 

symmicta (Lecanora) 30, 98, 206 

synothea (Lecidea) 128, 199 

synotheoides 24, 32, 57, 58*, 64, 74*, 88, 90, 91, 

93-95, 100, 102, 157, 180, 188-190* 

tenebrosa (Schaereria) 88 

ternaria 22, 23, 29, 59*, 63, 64, 83*, 89, 95, 96, 

99, 104, 126, 144, 171, 190, 191*, 192 

terrestris (Lecidea) 199 

tetramera (Bilimbia) 112 

Toninia 22 

trachona (Bacidia) 67, 206 

Trapelia 199 

Trapeliaceae 199 

Trapeliopsis 199 

Tremolecia 97 

trisepta 170,171,200 

triseptataNyl. 143 

triseptata Hepp (Bacidia) 143 

triseptatula 170 

triseptatuloides 170 

tuberculata 24, 25, 29, 31, 59*, 61, 62*, 63, 64, 

82*, 84, 87, 89, 90, 95, 100, 107, 163, 168, 187, 

192, 193*, 194, 204-206 

turfosa Massal. 24, 29, 32, 60*, 62, 64, 88, 89, 95, 

96, 103, 106, 155, 194, 195*, 196, 199, 204-206 

turfosa Fr. 47*, 150 

turgidula (Lecidea) 30, 168, 204-206 

Tylothallia 97 

uliginosa (Lecidea) 169, 206 

umbrinum (Scoliciosporum, Bacidia, 

,

214 

umbrinutn - cont. 

Lecidea) 25, 92, 97, 127, 128, 138, 140, 205, 

206 

umbrosa 149 

vainioi 186 

vermicillifera (Opegrapha) 

vernalis (Lecidea, Biatora) 

verrucula 194 

vezdae (Bacidia) 196 

Vezdaea 24,30,98 

67 

20, 205 


vinosa (Arthonia) 204 

violacea 51*, 170 

violacea (Bacidia) 170 

viridescens (Lecidea, Micarea) 

viridis (Protococcus) 174 

vulgaris 128 

vulgata (Opegrapha) 67 

xylophila 143 

zsakii 22, 48*, 154 

\ 

198,200,204,205

Bulletin of the British Museum (Natural History)

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