Foliicolous lichens from Madeira, with the description of a new ...

Lichenologist 28(3): 197–227 (1996) 

FOLIICOLOUS LICHENS FROM MADEIRA, WITH 

THE DESCRIPTION OF A NEW GENUS AND TWO 

NEW SPECIES AND A WORLD-WIDE KEY OF 

FOLIICOLOUS FELLHANERA 

Emmanuël SÉRUSIAUX* 

Abstract: A preliminary annotated list of the foliicolous lichens and of their 

lichenicolous fungi occurring in the laurisilva in Madeira is presented. Fellhaneropsis 

Sérus. & Coppins gen. nov. is introduced to accommodate Bacidia myrtillicola 

Erichsen and Bacidia vezdae Coppins & James, both being present in Madeira on 

living leaves. Two new species are described: Byssoloma kalbii Sérus. sp. nov. and 

Fellhanera seroexspectata Sérus. sp. nov. The following combinations are introduced: 

Fellhanera lambinonii (Sérus.) Lücking & Sérus., Fellhaneropsis myrtillicola (Erichsen) 

Sérus. & Coppins, F. vezdae (Coppins & James) Sérus. & Coppins, Woessia apiahica 

(Müll. Arg.) Sérus., W. canariensis (Lumbsch & Vězda) Sérus. and W. vasakii 

(Vězda) Sérus. A key to all foliicolous species of Fellhanera is presented. Bacidia buxi 

Vězda & Vivant and Bacidia gorgonea Vězda & Poelt are reduced into synonymy 

with Fellhaneropsis myrtillicola, Tapellaria similis Kalb with Byssoloma marginatum 

(Arnold) Sérus., and Bacidia michaeliana Sérus. with Bacidia lambinonii Sérus. 

1996 The British Lichen Society 

Introduction 

Madeira is a volcanic island in the North Atlantic Ocean, 565 km W of the 

African coast of Morocco and 3245N, 1700W. It is a serrated mountain 

range reaching 1861 m a.s.l. at the Pico Ruivo, and many deep rugged ravines 

run to the coast. Madeira belongs to the ‘macaronesian’ phytogeographical 

zone, together with the Canary Islands and the Azores, well known for its high 

level of endemism in flowering plants and for the persistence of an evergreen 

subtropical cloud forest, usually known as the laurisilva. In Madeira, this 

forest is best developed at 700–1200 m on the southern flank of the island and 

between 300 and 1300 m on the northern flank, if one includes the bushes 

dominated by Erica arborea and Myrica faya that top it on its higher parts. The 

tree and shrub flora (nomenclature following Hansen & Sunding 1993) is 

dominated by four species of Lauraceae (Appolonias barbujana (Cav.) Bornm., 

Laurus azorica (Seub.) Franco, Ocotea foetens (Ait.) Baill., and Persea indica 

(L.) K. Spreng.), together with Clethra arborea Ait. (Clethraceae), Myrica faya 

Ait. (Myricaceae), Ilex canariensis Poir. (Aquifoliaceae), Picconia excelsa (Ait.) 

DC. (Oleaceae), Heberdenia excelsa (Ait.) Banks ex DC. (Myrsinaceae), 

Euphorbia mellifera Ait. (Euphorbiaceae), Isoplexis sceptrum (L. fil.) Loud. 

(Scrophulariaceae), Erica arborea L. (Ericaceae), Vaccinium padifolium J. E. 

Sm. ex Rees (Ericaceae) and several others. The ground layer is extremely rich 

in ferns, including several endemic species. The laurisilva represents the 

*Research Associate F.N.R.S., Department of Botany, University of Liège, Sart Tilman, B–4000 

Liège, Belgium. 

0024–2829/96/030197+31 $18.00/0 1996 The British Lichen Society

198 THE LICHENOLOGIST Vol. 28 

Miocene forests of southern Europe, which vanished during the glacial times 

of the Pleistocene period, and is therefore of tremendous ecogeographical 

interest. 

Knowledge of the lichen flora of the island, as well as of the other islands of 

the archipelago, especially Porto Santo, has made great progress during recent 

years, the last contribution being that of Kalb & Hafellner (1992). That flora 

includes several endemics, e.g. Nephroma areolatum P. James & F. J. White for 

the main island, and Anzia centrifuga Haugan and several species of Ramalina 

for Porto Santo. A checklist has been prepared by Hafellner (1992) and is of 

great help for any study of the lichen flora of the archipelago. 

Foliicolous lichens have been reported from the island (see Santesson 1952) 

and, in the framework of a thorough study of this ecological group in western 

Europe (Sérusiaux 1993), I visited the laurisilva in February 1988 and in May 

1992 to gather comprehensive collections of these species. This paper deals 

with the first results of the study, and also includes collections made on 

the island by fellow botanists, especially L. Arvidsson, M. S. Christiansen, 

J. Duvigneaud, G. Een, J. Etayo, K. Kalb, H. Persson, S. Schuhmacher, C. 

Tavares and L. Tibell. This paper must, however, be considered as preliminary, 

as several taxa (at least three lichens and one lichenicolous fungus) have 

not yet been studied in detail and are not included in the list presented below. 

The study led to the reassessment of the generic position of Bacidia myrtillicola 

and of Bacidia vezdae, two species previously studied by my colleague and 

friend B. J. Coppins. He is therefore associated with the new genus described 

for them and to the two new combinations. 

The laurisilva has suffered much damage from human activities since the 

discovery of the island early in the fifteenth century and is now very reduced 

and severely degraded. The best remnants were explored, the localities 

visited being: Ribeiro Frio, along the Levada do Furado, 800–850 m alt.; 

Portela, along the Levada de Portela, near Lamaceiros, 800 m alt.; Casa de 

Queimadas, track towards Caldeirâo Verde, 850 m alt.; Châo do Louros, N of 

Encumeada pass, 800 m alt.; the Riba da Seixal, S of Seixal at 300–400 m alt. 

Foliicolous lichens have been found on the following tree or shrub species: 

Appolonias barbujana, Laurus azorica, Persea indica, Ocotea foetens, Picconia 

excelsa, Ilex canariensis, Heberdenia excelsa, Clethra arborea, and on the following 

herbaceous plants growing at ground level: Ruscus streptophyllus P. F. Yeo 

(Liliaceae) and Trichomanes speciosum Willd. (Hymenophyllaceae). Several 

introduced species also have foliicolous species, e.g. Camellia japonica 

(Theaceae), when they are planted in suitable localities. 

The Genus 

Fellhaneropsis Sérus. & Coppins gen. nov. 

Thallus crustaceus, tenuis, sine cortice, et algas verosimiliter Chlorococcaceae continens. 

Ascomata apothecia, 0·1–0·4 mm diam., basi arcte constricta, cum margine tenui mox evanescenti. 

Excipulum tenue sed proprium, cum ellipticis et polyedralibus cellulis, in verticalibus 

seriebus dispositis, typicam paraplectenchymaticam texturam non formantibus; hamathecium 

cum ramosis et anastomosantibus paraphysibus; asci ad Byssoloma typum (sensu Hafellner 1984: 

315) pertinentes, octospori; sporae oblongae-fusiformes ad fere aciculares, cum transversalibus 

septis. Pycnidia longa filiformiaque conidia producentia, et (solum in Bacidia myrtillicola) parva 

bacilliformiaque. 

Type: Bacidia myrtillicola Erichsen

1996 Foliicolous lichens from Madeira—Sérusiaux 199 

Thallus crustose, thin, without cortex and with most probably a species of 

Chlorococcaceae as photobiont. Ascomata apothecia, 0·1–0·4 mm diam., 

distinctly constricted at their base, with a thin, soon disappearing margin. 

Excipulum thin but distinct, with elliptical to polyhedral cells, arranged in 

upright rows, not forming a typical paraplectenchymatous tissue. Hamathecium 

of branched and anastomosed paraphyses. Asci of the Byssoloma-type 

(sensu Hafellner 1984: 315), 8-spored. Ascospores oblong-fusiform to almost 

acicular, with transverse septa. Pycnidia producing long and filiform conidia, 

and in addition (in Bacidia myrtillicola only) smaller bacilliform conidia. 

Bacidia myrtillicola and Bacidia vezdae are obviously close to the genus 

Fellhanera Vězda in the Pilocarpaceae (Vainio) Zahlbr. (Vězda 1986). Both 

species here transferred to the new genus Fellhaneropsis share all typical 

characters of the Pilocarpaceae and could be referred to Fellhanera, as 

accepted by several authors. We consider, however, that their excipulum type 

(elliptical to polyhedral lumina, arranged in rows) and their filiformsigmoid 

conidia are good apomorphies that require the recognition of a 

different genus. Fellhanera s.str. includes species with a typically paraplectenchymatous 

excipulum (isodiametric cells), and bacilliform to ellipsoid, or 

pyriform-clavate conidia. 

Most species of Fellhanera Vězda have pyriform-clavate conidia, as do the 

most common species in Europe (F. bouteillei (Desm.) Vězda and F. subtilis 

(Vězda) Diederich & Sérus.). The type species (F. fuscatula (Müll. Arg.) 

Vězda), however, has bacilliform to mostly ellipsoid conidia (a feature not 

quoted by Vězda 1986: 201). This type of conidium is also present in Bacidia 

myrtillicola (microconidia; see below), but this species cannot be considered as 

congeneric with F. fuscatula as the latter has a typically paraplectenchymatous 

excipulum and never produces filiform conidia (numerous collections seen, 

mainly from tropical Africa, LG). Indeed, the filiform-sigmoid conidia 

produced by both species here referred to the new genus are probably the best 

apomorphy for it, as well as the apothecial origin of the pycnidia producing 

them. This is clearly demonstrated for Fellhaneropsis myrtillicola in this paper 

and is strongly suspected for F. vezdae. This feature appears to be unique in 

lichenized fungi and clearly merit the generic status within the Pilocarpaceae. 

The genus Badimia Vězda, so far assigned to the same family, produces highly 

specialized pycnidia (campylidia) with extraordinary conidia (very long and 

filiform, multi-septate, with crooked ends and lateral appendages terminated 

by a tiny, sticky mass) (Sérusiaux 1986). The primordia of these campylidia 

are impossible to distinguish from those of apothecia and their ontogeny 

shows that they are transformed apothecia. 

The Species 

Fellhaneropsis myrtillicola (Erichsen) Sérus. & Coppins comb. nov. 

Bacidia myrtillicola Erichsen, Mitt. Inst. Allg. Bot. Hamb. 10: 414 (1939). Fellhanera myrtillicola 

(Erichsen) Hafellner, In Vězda, Lich. Sel. Exs. 95, No. 2358 (1989); type: Germany, Schleswig- 

Holstein, Lauenburg, im Sachsenwald bei Friedrichsruh, an Vaccinium myrtillus am Rande des 

Reviers Saupark, 2 xi 1924, C. F. E. Erichsen (HBG—holotypus!).


Bacidia myriocarpa Erichsen, Mitt. Inst. Allg. Bot. Hamb. 10: 413 (1939); type: Germany, 

Schleswig-Holstein, Niedersachen, Kreis Lüneburg, Findling im Frost Einemhof, bei Radburch, 

14 iii 1937, Lehr (HBG—lectotypus!, selected by Jacobsen & Coppins 1989: 259). 

Bacidia nitschkeana var. perpusilloides Erichsen, Ann. Mycol. 41: 206 (1943); type: Germany, 

Schleswig-Holstein, Flenburg, Forst Clusries, near Wasserleben, on Picea twigs, 6 x 1923, C. F. 

E. Erichsen (HBG—holotypus!). 

Bacidia buxi Vězda & Vivant, Bull. Soc. Bot. Fr. 119: 256 (1972). Fellhanera buxi (Vězda & 

Vivant) Vězda, Folia Geobot. Phytotax. (Praha) 21: 214 (1986); type: France, Dépt. Pyrénées- 

Atlantiques, Sauveterre-de-Béarn, île sur le gave d’Oléron, sur feuilles de Buxus sempervirens, 

28 i 1972, Vivant (hb. Vězda-holotypus!; UPS—isotypus!, isotypi also distributed in Vězda Lich. 

Sel. Exs. No. 1161, BM, G, LG!). 

Bacidia gorgonea Vězda & Poelt, in Poelt & Vězda, Herzogia 9: 241 (1992); type: Österreich, 

Steiermak, Windische Bühel, enger W-E-verlaufender westlicher Seitengraben des Gamlitzbach- 

Tales, S Kranach, WSW Gamlitz, auf Nadeln von Abies alba, 6 x 1991, Poelt & Giralt 

(GZU—lectotypus, selected here, see below!). 

(Figs 1–2) 

Illustrations: Vězda & Vivant (1972: 257); Coppins (1983: 197, fig. 56A); Poelt & Vězda (1992: 

245) and Arup & Ekman (1994: 37, fig. 3C & 39, fig. 4C). 

Thallus formed of small roundish patches that eventually merge to form 

a continuous thallus up to several centimetres in diam., thin and smooth 

or slightly verrucose, or convex and formed of agglutinated granules, 

especially when the thallus grows along the main vein of a leaf, or near the 

nodes of a twig or on a rough surface (rock or bark), greenish grey to bluish 

grey (the typical specimens have a distinct bluish tinge), usually with a bluish 

pellucid prothallus, best seen on foliicolous specimens, with a shiny green to 

dark green surface colour when invaded by epiphytic green algae and 

cyanobacteria. Photobiont: most probably a species of Chlorococcaceae, with 

spherical, green cells, 5–12 μm diam. Thallus reactions: K, C,P. 

Apothecia usually numerous, sometimes absent, very rarely agglutinated and 

never proliferating, circular or rarely with a swollen or irregular edge, distinctly 

constricted at the base, 0·1–0·2 mm diam., mostly not exceeding 0·15 mm, 

exceptionally up to 0·3 mm diam., less than 0·1 mm in height, at first flat 

and with a thin and pale but nevertheless distinct margin, soon convex and 

immarginate; disc pale brown to bluish grey, sometimes almost mauvish 

(some specimens show a patchwork of pale brown and bluish grey apothecia). 

Excipulum distinct but not exceeding 15 μm in thickness, with elliptical to 

polyhedral lumina, arranged in upright rows, never forming a typical 

paraplectenchymatous tissue (isodiametric lumina), hyaline but with a brownish 

to greenish colour in the zone adjacent to the hymenium. Hypothecium less 

than 10 μm thick, dark brown, K+ greenish brown to aeruginose green, 

usually with the K+ reaction restricted to the central and basal parts. 

Hamathecium of branched and anastomosed paraphyses, 1–1·5 μm thick, 

much branched in the upper parts to form a densely interwoven network 

around the ascus tops and in the epithecium. Asci of the Byssoloma-type 

(sensu Hafellner 1984: 315), 30–4012–16 μm, 8-spored. Ascospores 

oblong-fusiform, straight or slightly curved, sometimes deformed, rarely 

tapering towards one end, sometimes with a thin perispore, 3(–5)-septate, 

16–25(–28)3–4 μm (one spore seen reached 314 μm), several times seen 

with germinating tubes at one end while in the asci.


FIG. 1. Fellhaneropsis myrtillicola (Spier W5030), pycnidia producing long and filiform conidia, at 

their early stage of development in spore-producing apothecia. A, Marginal section; B, Radial 

section, a fully-developed ascus is clearly seen in the centre. Scale=0·1 mm.


FIG. 2. Fellhaneropsis myrtillicola (Spier W5030), pycnidium producing long and filiform conidia, 

at its early stage of development in a spore-producing apothecium. Radial section; several asci are 

still producing ascospores. Scale=0·1 mm. 

Pycnidia of two types: (i) usually present, producing long and filiform 

conidia (macroconidia) and (ii) abundant or totally absent, producing small 

bacilliform conidia (microconidia). 

(i) Conidiogenous layer first appearing as a circlet deep in spore-producing 

apothecia, in the basal zone between the hypothecium and the excipulum, 

rapidly expanding and producing conidia, and ejecting the hymenium and 

hamathecium, which degenerate and finally disappear; some time after the 

appearance of the conidia, the upper part of the excipulum (=apothecial 

margin) starts growing upwards long, thin-walled hyphae, gently tapering 

towards their top and thus forming a vertical to oblique cylinder of ‘palisadic’ 

hyphae with a fimbriate rim at the top, and containing the conidia; mature 

pycnidia 0·1–0·15 mm diam. and up to 0·2 mm in height, but usually not 

exceeding 0·1 mm; in the best developed ones, the conidial mass is seen as a 

translucent ball under the dissecting microscope, whereas in eroded or in 

dying pycnidia, conidia and/or the ‘palisadic’ hyphae can be totally absent; 

conidiogenous cells elongate, 10–203–5 μm. Conidia long, filiform-sigmoid, 

non-septate, sometimes irregularly twisted and with a swollen end, 20–45 

1–1·5 μm. 

(ii) Sessile and globose, sometimes deformed, usually constricted at their 

base, 0·05–0·08 mm diam., sometimes with a glut of conidia at their apical


ostiole; wall green-brown to bluish green; conidiogenous cells carpeting the 

base of the pycnidium, elongated-polyhedral. Conidia rare or numerous, 

bacilliform to cylindrical, sometimes narrowly ellipsoid, often deformed (with 

swollen parts, irregularly bent), 4–80·5–1 μm (in some specimens, conidia 

reaching 10–131 μm are present, mixed with ‘normal’ ones). 

Notes: Fellhaneropsis myrtillicola is usually difficult to detect on any substratum 

on which it can grow: it rarely forms a dominant feature of any 

community, its thallus is easily passed over as a dying crust and its apothecia 

are tiny and without any brightness of colour. The best criterion to identify it 

is to search for the pycnidia producing macroconidia. They are usually present 

and highly characteristic: the ‘palisadic’ hyphae in the pycnidia are, to our 

knowledge, unique for European and Macaronesian lichens. 

Long, filiform-sigmoid conidia are not common in crustose lichens; there is 

a risk of confusion with the Micarea peliocarpa-group (Coppins 1983: 99 & 

112), which also produce long and curved conidia and which can also have 

pale bluish apothecia with 3-septate ascospores. They are easily distinguished 

by their apothecium size (mostly over 0·2 mm diam., and reaching at least 

0·4 mm diam.; such sizes are never reached in F. myrtillicola), by their 

excipulum (made of richly branched and anastomosing, radiating hyphae, 

never with elliptical-polyhedral lumina as in F. myrtillicola), by their thallus 

and apothecia reacting C+ red (production of gyrophoric acid, which is absent 

in F. myrtillicola) and by their pycnidia, not arising from apothecia and devoid 

of any ‘palisadic’ hyphae. 

The species can also be confused with immature or foliicolous forms of 

Byssoloma marginatum (Arnold) Sérus.; the differences between these species 

are provided in the following paragraph, under the analysis of the epithet 

gorgonea. 

Notes on the Use of the Epithets 

—myrtillicola. This epithet was introduced simultaneously with myriocarpa 

by Erichsen and was adopted for the species dealt with here by Jacobsen & 

Coppins (1989: 258) on the basis of Art. 57.2 of the ICBN. It was erroneously 

used by Hafellner (in Vězda Lich. Sel. Exs. No. 2358 & 2359, 1989) and by 

Poelt & Vězda (1990: 389) for Fellhanera subtilis (Vězda) Diederich & Sérus. 

This error has been detected by Diederich et al. (1991: 3) and by Arup & 

Ekman (1994: 37–38). The type specimen of Bacidia myrtillicola has already 

been studied by Coppins (1983: 196). 

This epithet has been scarcely used in the literature: apart from the 

examination of the Erichsen’s original collections by Coppins (1983: 196) and 

by Jacobsen & Coppins (1989: 258–259), we have noted the records by 

Benfield (1992: 42; specimen checked), Jacobsen (1992: 78; one specimen 

seen), and by Arup & Ekman (1994; specimens examined). 

—myriocarpa. I had the opportunity to study the type collection as well as 

the other specimen mentioned by Jacobsen & Coppins (1989: 259): ‘Same 

locality [as the lectotype], 12 iv 1937, C. F. E. Erichsen’ (HBG). Bacidia 

myriocarpa Vězda (Vězda 1975a: 123–124) is a later homonym, and is now 

regarded as a synonym of Fellhanera rhapidophylli (Rehm) Vězda.


—perpusilloides. B. J. Coppins (Coppins 1983: 196) has examined the 

lectotype and paratypes. A further specimen annotated Bacidia nitschkeana 

var. perpusilloides by Erichsen (‘Kreis Lanenburg, an jungen Fichtenzweigen 

im Sachsenwald, 11 v 1924, C. F. E. Erichsen’, HBG) has also been examined: 

it contains only Fellhanera subtilis and Scoliciosporum chlorococcum (Graewe ex 

Stenh.) Vězda. 

—buxi. The type collection is plentiful and I have been able to examine 

several parts of it; moreover I visited the locality in 1985 and again in 1989, 

together with P. W. James, F. Rose and J. Vivant. This population shows all 

the characters of Fellhaneropsis myrtillicola: thallus, apothecia and both types of 

pycnidia. Some isotypes distributed in the Vězda Lich. Sel. Exs. may be 

disconcerting as they exuberantly produce apothecia that are pale brown and 

lack the typical bluish tinge of the species; moreover in such specimens, the 

pycnidia with filiform conidia are absent. Otherwise, the collections gathered 

in this locality are typical. 

A great deal of confusion is present in the literature concerning the use of 

this epithet: the specimens mentioned under this name from Tenerife 

(Lumbsch & Vězda 1992: 25) belong to Fellhanera christiansenii Sérus. & 

Vězda (see below); those mentioned from S Austria (Poelt & Vězda 1992: 243; 

specimens seen, GZU!) are typical Fellhanera subtilis, and those reported from 

the western Caucasus (Vězda 1983: 63; specimens seen, LG and hb. Vězda!) 

belong to a further, still undescribed species of Fellhanera. Indeed, these 

Caucasian specimens have a typical paraplectenchymatous excipulum and 

pyriform to obclavate conidia. E. Sérusiaux will deal with this taxon as soon as 

its relationships with a foliicolous population of a puzzling Fellhanera from 

Spain/Cataluña are solved. The material mentioned from Costa Rica by 

Lücking (1992: 136; Costa Rica, Lücking 88-158, hb. Lücking!) is too scanty 

for a definite identification but there is no doubt it is not Fellhaneropsis 

myrtillicola. Fellhanera buxi has also been reported in very small quantity on 

Calluna stems in Ireland (McCarthy et al. 1985: 96) but the material received 

on loan (1978, M. G. C. Schouten, GALW!) does not contain this species. 

Associate foliicolous species at the type locality include: Woessia chloroticula 

(Nyl.) comb. ined. (=Bacidia chloroticula (Nyl.) A. L. Sm.), Byssoloma 

leucoblepharum (Nyl.) Vainio, Fellhanera bouteillei (Desm.) Vězda with 

Wentiomyces lichenicola (Hansf.) D. Hawksw. subsp. bouteillei Bricaud et al. 

(Roux et al. 1994: 473) growing on its thallus, Porina hoehneliana (Jaap) R. 

Sant., P. leptosperma Müll. Arg. and P. oxneri R. Sant. Phorophytes are 

cladodes of Ruscus aculeatus, leaves of Buxus sempervirens and of introduced 

species of Elaeagnus and of Diospyros. Fellhaneropsis myrtillicola grows on leaves 

of Buxus and of Elaeagnus. 

—gorgonea. The type collection (GZU!), which is plentiful and covers rather 

large parts of needles and twigs of Abies, is a mixture of Byssoloma marginatum 

(Arnold) Sérus. and of Fellhaneropsis myrtillicola. The original description 

(Poelt & Vězda 1992: 241–242) is based on the Byssoloma for the apothecia 

and on the Fellhaneropsis for the pycnidia. The only pycnidia present (with 

macroconidia) are very typical of F. myrtillicola and show almost all stages 

between immature ones and ‘fully-mature’ ones with a glut of macroconidia


raised between the typical walls of the species’ pycnidia, including the circlet 

of elongate thin-walled hyphae (named ‘Palisaden-Hyphen’ by Poelt & Vězda; 

their fig. 1, pv is an old, collapsing pycnidium). 

Two different types of apothecia are present: those of F. myrtillicola, 

distinctly constricted at their base, with almost immarginate, convex, pale 

bluish and sometimes pale brown discs, and those of B. marginatum, almost 

sessile, with flat, clearly marginate (swollen, livid pale bluish grey), and 

bluish grey discs. Sections of the apothecia of both species are very similar, 

especially as the asci both belong to the Byssoloma-type (sensu Hafellner 1984: 

315) and as the hypothecium pigment is closely related (K+ greenish brown 

to green-aeruginose in F. myrtillicola and K+ dark aeruginose, or greenish 

brown to purple-brown in B. marginatum). Only a detailed study can give 

prominence to the differences: in F. myrtillicola, the excipulum is made of 

hyphae with elliptical to polyhedral lumina, arranged in upright rows (in 

squash preparations, it is usually possible to see fragments that can be 

described as paraplectenchymatous) while in B. marginatum, the excipulum is 

made of tightly interwoven hyphae that separate with difficulty in a KOH 

solution. In the type collection, the original authors have separated an 

Abies needle with typical apothecia of B. marginatum in a small envelope, 

labelled as ‘Holo’. They describe the excipulum as ‘prosoplektenchymatisch, 

die strahlig angeordneten Hyphen dicht meist ungeteilt’, which clearly points 

to B. marginatum, and their description of the ascospores (‘oft schlecht 

entwickelt, ellipsoid oder ellipsoid-spindelig, oft an einem Ende verschmälert, 

3-, selten 1-septiert, 14–183–3·5 μm’) is a perfect one for young stages of B. 

marginatum. Indeed, this usually corticolous species normally produces rather 

large apothecia (up to 0·5–0·7 mm diam.) with ellipsoid, straight or hardly 

curved, 3-septate ascospores, measuring when fully mature 20–224–5 μm. 

However, the species is also foliicolous (especially in Madeira) and then 

rapidly produces apothecia with ascospores that are 1–3-septate, curved, 

attenuated at one end and measure 12–183–4 μm. Field experience of that 

species in Madeira shows that this is only an immature stage. The ascospores 

of F. myrtillicola very quickly reach their final septation and dimensions, and 

are oblong-fusiform, straight or slightly curved, rarely tapering towards one 

end, 3(–5)-septate, 16–25(–28)3–4 μm; usually there is no problem in 

distinguishing between both species on the basis of mature ascospores. 

Poelt & Vězda (1992) obviously based their new species on the pycnidia 

producing long and filiform conidia. In spite of their marking of an Abies 

needle with apothecia of Byssoloma marginatum as the holotype, we lectotypify 

the epithet gorgonea on the apothecia and pycnidia of the species named 

Fellhaneropsis myrtillicola in this paper. This decision is in accordance with art. 

9.10 of ICBN. However, we could have decided to follow Rec. 9 A.3 of ICBN 

and lectotypify the epithet on the material marked as ‘holotype’; in this case, 

Bacidia gorgonea would have become a synonym of Byssoloma marginatum 

(Arnold) Sérus. 

Pending a final decision on the status of myrtillicola, buxi and gorgonea, E. 

Sérusiaux has used this latter epithet to name specimens from southern France 

submitted to him by Dr C. Roux. They have been published under that name 

(Bricaud et al. 1993: 100).


—Lecidea quintula Ny., Flora 48: 5 (1865). Type: France, Brest, on stem of 

Calluna, Crouan (H-NYL 19061, 19062 & 19063—syntypi!). 

Dr C. Printzen kindly drew my attention to this forgotten name and 

arranged for the loan of the syntypes preserved in H. All syntypes contain 

fragments of a species that might be Fellhaneropsis myrtillicola. A short 

description follows: thallus thin, greyish; apothecia 0·16–0·35(–0·4) mm 

diam., convex and without margin, dark brown to black; excipulum of 

outwardly radiating hyphae with polyhedral lumina; hypothecium brown, 

with a K+ olivaceous reaction; hamathecium of branched and anastomosed 

paraphyses; asci of the Byssoloma-type; ascospores oblong-fusiform, 

3–5-septate, with a thin perispore, 20–263–4(–4·5) μm, some with a 

germinating tube. 

This description clearly points to Fellhaneropsis myrtillicola, except for a few 

details regarding the apothecia (size and colour). I do not know of any other 

European species with Byssoloma-type asci and such long ascospores to which 

these collections may be referred, and their ecology (epiphytic on Calluna 

stem) is compatible with that of F. myrtillicola (see below). I have, however, 

refrained from using this epithet (which is much older than myrtillicola) for the 

following reasons: the type material is very scanty and obviously badly 

preserved, and does not allow a thorough investigation; no pycnidia can be 

found on them (macroconidia are definitely the best criterion to identify 

F. myrtillicola); the colour of the apothecia and of the thallus are not typical of 

the species (but these features might be explained by the bad preservation of 

the material); the apothecia size is abnormal for the species (but this may be 

explained by its growing on a much more stable substratum than twigs and 

leaves); and the flora of Calluna stems in Brittany is very poorly documented 

and may yield further, little studied species. I have, therefore, decided to wait 

until fresh data on this flora are available to decide upon the status of Lecidea 

quintula Nyl. 

The syntype H-NYL 19061 also contains fragments of Byssoloma subdiscordans, 

and H-NYL 19062 has poor but typical (apothecia and pycnidia) 

B. marginatum. As the protologue says that the ascospores are 5-septate and 

measure 21–255–6 μm, it is clear that Nylander was not dealing with these 

two species in describing Lecidea quintula. 

Fellhaneropsis myrtillicola, as circumscribed in this paper, is a widespread but 

overlooked species. It is now known from western and central Europe, and 

from Macaronesia. I expect that more attention to this tiny species will 

demonstrate that it is not rare in suitable habitats. As is the case with several 

other ‘foliicolous’ lichens, such as Fellhanera bouteillei, Fellhaneropsis myrtillicola 

is a weak competitor but is able to proliferate on quite different substrata in 

very different habitats, including semi-natural disturbed ones. 

It is known on acidic rocks in W England and N Germany (Jacobsen 1992: 

78), in shaded and humid environments. It grows on Vaccinium twigs in dense 

thickets at forest margins or inside forests in S Sweden (see Arup & Ekman 

1994 for more details) and in the Belgian Ardennes. In central Germany, it has 

recently been found in Calluna twigs in a cemetery, which clearly shows that 

the species is able to invade highly artificial habitats. It is also present on Abies


twigs in hilly or mountainous areas of Germany, Austria and of Calabria in 

southern Italy; its ecology in such habitats is thoroughly described by Poelt & 

Vězda (1992). It is also present on Buxus leaves and twigs in S Belgium, in 

southern (s. lat.) France and in the eastern parts of the Spanish Pyrenees. In 

Belgium, the locality is the last remnants of Buxus shrubs at the bottom of a 

humid, shaded valley (a more detailed account of this fascinating site is in 

preparation by P. van den Boom and E. Sérusiaux), while in France and in 

Spain, it tolerates drier and fluctuating conditions in the summer but 

nevertheless requires a very humid atmosphere at certain periods of the year, 

at least during winter (O. Bricaud is studying the ecological conditions of the 

numerous sites with a foliicolous lichen flora in Provence/France). In the 

laurisilva of Tenerife and Madeira, it develops only small, dispersed and 

ill-looking thalli; it is obviously unable to compete with the most common 

foliicolous species (Byssoloma and Fellhanera species) and is sometimes the 

only species present, usually along the main vein, together with Gyalectidium 

colchicum, which grows mainly on the leaf surface. 

Additional specimens examined: Sweden: Västergötland, Kinnarumma par.: Flenstorp, mixed 

coniferous-deciduous forest, on twigs of Vaccinium myrtillus, viii 1993, Arup & Ekman (LD). 

Småland, Annerstad par.: V. Ringabergsholmen, small islet of damp Picea-Populus forest in a large 

bog, on twigs of V. myrtillus, vi 1993, Arup & Ekman (LD); Hemmesjö par.: c. 200 m SSW of 

Vitarör, on small twigs of Frangula alnus in a rather old Picea forest, vii 1993, Arup (LD); 650 m 

NW of Fridhem, E of Store mosse, old coniferous forest, on twigs of V. myrtillus, iii 1994, Arup 

(LD); c. 500 m S of Vitarör, rather old Picea forest, on twigs of V. myrtillus, vi 1993, Arup (LD). 

Skåne, Örkened par.: Nytebodaskogen, at Lommagylet, old Picea forest, on twigs of V. myrtillus, 

xii 1993, Ekman (LD).—Germany: See type collections Bacidia myriocarpa and of Bacidia 

nitschkeana var. perpusilloides. ‘An Zweigen junger Fichten am Nordosthange der Kalkenberges, 

bei Klangenfurt’, s.d., J. Steiner (B); another specimen is preserved in B but the label is illegible. 

Feldweg zwischen Emkendort und Brux, an Steinen auf einem flachen, erdbebedeckten Blockwall 

am Waldrand sowie an Steinem unmittlebar auf den Waldboden, iv 1989, Jacobsen 6122 (E). 

Nord-Niedersaschen, LK Harburg, cemetery in Lübberstedt, on Calluna twigs, ix 1994, G. Ernst 

s.n. (E).—Great Britain: N Devon, v.c.4: Oakford, near Spurway Baston, Combe Water, on top 

of a stream boulder, 1992, Benfield s.n. (E).—Netherlands: Den Treek, 4 km SE of Amersfoort, 

on Vaccinium myrtillus, ii 1994, Spier W5030 (hb. Spier).—Belgium: Namur prov.: Waulsort, 

Fonds des Vaux, small wooded valley with mature Buxus sempervirens, on twigs and leaves of 

Buxus, 14 iv 1993, van den Boom 13871 (hb. van den Boom); ibid., vi 1994, Sérusiaux s.n., 

Diederich 4996 & van den Boom (LG, hb. Diederich). Luxembourg prov.: Anlier, rivulet ‘Fond du 

Gris Boffet’, shrubs of Calluna vulgaris and of Vaccinium myrtillus by a disused meadow, on twigs 

of Calluna and Vaccinium, x 1988, Sérusiaux 10246a (LG).—Austria: see type collection of 

Bacidia gorgonea.—France: see type collection of Bacidia buxi. Dépt. Isère: Cognin-les-Gorges, 

Gorges of Nant, dense wood with Buxus sempervirens and Corylus avellana, on leaves of Buxus, 

viii 1986, Sérusiaux s.n. (LG). Dépt. Aveyron: Gorges of the Lot, along the road to Issac and 

Florentin-la-Chapelle, dense wood with B. sempervirens, on leaves of Buxus, viii 1986, Sérusiaux 

s.n. (LG). Dépt. Pyrénées-Occidentales: Sauveterre-le-Béarn, île sur le Gave d’Oloron, disturbed 

forest with mature B. sempervirens, on leaves of Buxus, vii 1985, Sérusiaux s.n. (LG); ibid., on 

leaves of Buxus and of Elaeagnus, vii 1989, Sérusiaux s.n. with James, Rose & Vivant (LG) (type 

locality of Bacidia buxi); Ste-Engrâce, Gorges of Ujarre, mixed vegetation with B. sempervirens and 

Corylus avellana, on leaves of Buxus, 18 vii 1991, Diederich & Etayo (LG). Dépt. Vaucluse: 

Méthamis, ‘vallon Peynier’, dense shrubs of B. sempervirens in a disturbed wood, on leaves of 

Buxus 3 v 1989, Roux s.n. with Bricaud & Clauzade (MARSSJ, LG); ibid., vii 1993, Sérusiaux s.n. 

with Bricaud & Roux (LG); Petit Lubéron, Ménherbes, small valley E of Mau Vallon, dense 

shrubs of B. sempervirens in a Quercus pubescens wood, on leaves of Buxus, vii 1993, Sérusiaux s.n. 

with Bricaud & Roux (LG); Petit Lubéron, Sivergues, Chantebelle, ‘vallon de l’enfer’, dense 

shrubs of B. sempervirens, on leaves of Buxus, vii 1993, Sérusiaux s.n. with Bricaud & Roux 

(LG).—Spain: Catalyuña, Girona prov.: Oïx, Riera d’Oïx, forest with B. sempervirens and Quercus 

pubescens, on leaves of Buxus, ii 1991, Etayo (hb. Etayo, LG).—Italy: Calabria: Vibo Valentia,


Serra San Bruno, nel Bosco di Santa Maria, on needles of Abies, iv 1993 and vi 1993, Puntillo s.n. 

(LG).—Madeira: Along the levada between Ribeiro Frio and Lombo Capitâo Mormo, on leaves 

of Lauraceae, vi 1952, H. Persson 98 (S); Ribeiro Frio, along the levada de Portela, disturbed 

laurisilva along the river, on leaves of Lauraceae and of an unknown introduced shrub, ii 1988, 

Sérusiaux s.n. (LG, 2 specimens); Châo do Louros, N of Encumeada pass, disturbed laurisilva, on 

leaves of Clethra arborea, ii 1988, Sérusiaux s.n. (LG); Châo do Louros, little disturbed laurisilva, 

on fronds of Trichomanes speciosum, v 1992, Sérusiaux s.n. (LG); Casa das Queimadas, track to 

Caldeirâo Verde, disturbed laurisilva, on Lauraceae, v 1992, Sérusiaux s.n. (LG); Riba do 

Seixal, S of Seixal, pristine laurisilva, on leaves of Lauraceae, v 1992, Sérusiaux s.n. (LG).— 

Canary Islands: Tenerife: Montaña de Anaga, near Pico del Inglés, laurisilva, on leaves of Ilex 

canariensis, xii 1978, Arvidsson s.n. (GB). 

Fellhaneropsis vezdae (Coppins & James) Sérus. & Coppins comb. 

nov. 

Bacidia vezdae Coppins & James, Lichenologist 10: 190 (1978). Fellhanera vezdae (Coppins & 

James) V. Wirth, Die Flechten Baden-Württembergs: 511 (1987). 

Type: Great Britain, England, West Sussex, Midhurst, ad corticem Querci, 8 December 1970, 

Coppins & Rose s.n. (BM—holotypus!). 

Illustrations: Coppins & James (1978: 192, fig. 2). 

A thorough description of this species is provided by Coppins & James 

(1978: 190–194). There is no doubt that this species is congeneric with 

myrtillicola as it has the same type of ascus structure, of ascospores and conidia. 

The main differences are the colour of its apothecia (livid brown to pinkish 

brown versus bluish to pale brown in myrtillicola) that frequently become 

tuberculate (very rarely in myrtillicola), the absence of any K reaction in the 

hypothecium (usually K+ greenish brown to green-aeruginose in myrtillicola), 

the septation and the size of its ascospores [(3–)5–7, 30–35(–42)3–4·5 μm 

versus 3(–5), 16–283–4 μm inmyrtillicola]. The relationships between both 

species were already highlighted by Coppins (1983: 196; under Bacidia 

nitschkeana var. perpusilloides). 

Although many collections were carefully examined, we are not able to 

demonstrate, as is the case in Fellhaneropsis myrtillicola, that the pycnidia arise 

within ascospore-producing apothecia and that the ‘excipulum’ elongate to 

form a circlet of raised hyphae around the ostiole. There are however clues 

that the pycnidia of F. vezdae derive from apothecia: the pycnidial wall has the 

same structure as the excipulum and the same pale brown K pigment, and 

well-preserved pycnidia have a ‘fimbriate rim’ (as already pointed by Coppins 

& James 1978: 191). 

Fellhaneropsis vezdae is known from western and central Europe (Purvis et al. 

1992: 112), usually growing on bark, and more rarely on bryophytes, other 

lichens or rock. It usually occurs in shaded habitats and tolerates a moderately 

polluted atmosphere. In Madeira, this species has been found several times in 

the laurisilva, but always in small quantities on living leaves, including 

Trichomanes speciosum fronds. It has just been reported from Madeira by Kalb 

& Hafellner (1992: 64) growing on branches of Erica arborea. 

Byssoloma kalbii Sérus. sp. nov. 

Foliicola Byssoloma species insignis apotheciis minutissimis et albidis, hypothecio hyalino et 

pycnidiis albidis.


FIG. 3. Byssoloma kalbii (holotype), thallus with apothecia and pycnidia on frond of Trichomanes 

speciosum. A, General view; B, Young apothecia; C, Ripe apothecium with marginal pycnidia. 

Scale=1 mm. 

Type: Portugal, Madeira, S of Seixal, Riba do Seixal, alt. 300–400 m, little disturbed laurisilva, 

on fronds of Trichomanes speciosum growing on the ground, May 1992, Sérusiaux s.n. (LG— 

holotypus).


Thallus epiphyllous, farinose but rather compact, with an irregular surface, 

matt, pale green with tinges of yellow or blue, continuous over the leaf 

surface. Photobiont: most probably a species of the Chlorococcaceae, with 

spherical, green cells, 5–12 μm diam. 

Apothecia usually numerous, sometimes absent, circular when young, 

eventually becoming lobulate, with an irregular edge, 0·15–0·2(–0·3) mm 

diam., less than 0·1 mm in height, without any distinct margin, disc very pale 

to pale brownish orange, almost white when young; becoming brownish when 

old, flat or convex. Excipulum poorly developed, best seen in young apothecia, 

almost absent in older ones, hyaline and without crystals, made of a loose 

network of branched and anastomosed hyphae, cells without any swelling at 

the margin. Hypothecium hyaline, less than 10 μm thick. Hamathecium of 

branched and anastomosed paraphyses. Asci of the Byssoloma-type (sensu 

Hafellner 1984: 315), 35–4510–15 μm, 8-spored. Ascospores ellipsoid, 

3-septate, not constricted at their septa, with a distinct gelatinous halo, 

10–142·5–4 μm. 

Pycnidia always numerous, sessile, globose, usually distinctly constricted at 

their base, 0·1–0·15 mm high, 0·05–0·1 mm diam., sometimes with a spherical 

glut of conidia at their apical ostiole, wall white to very pale brown. Conidia 

numerous, bifusiform to slightly clavate, 4–51–1·5 μm. 

Byssoloma kalbii belongs to a small group of foliicolous Byssoloma species 

with tiny apothecia almost devoid of a margin and with 3-septate ascospores. 

These species are weak competitors and are usually present in atypical niches, 

such as the leaf margins or the fronds of fragile ferns, or at early stages of 

the colonization of leaves by foliicolous organisms. Byssoloma kalbii is easily 

distinguished from related species by the very pale apothecia, hyaline 

hypothecium and the abundant, white globose pycnidia. 

The related species are (Vězda 1975b: 426–427, Kalb & Vězda 1990: 

445–447, Barillas & Lücking 1991: 314, Lücking 1992: 143): 

—Byssoloma subpolychromum Vězda, known from Africa/Tanzania and Zaïre 

and from ‘Amerika’ (fide Vězda 1987: 74, but not mentioned in Kalb & Vězda 

1990), and which has pale brown to brown apothecia, 0·3–0·5 mm diam. and 

a dark brown hypothecium. Pycnidia are also abundant and have a dark brown 

wall. 

—Byssoloma minutissimum Kalb & Vězda, widespread in Brazil, also found 

in Guatemala and Costa Rica, and which has pale ochre apothecia, 0·15– 

0·2 mm diam. and a brown hypothecium. Pycnidia have not yet been found in 

this species. 

Further data about species of Byssoloma with tiny apothecia and hardly 

distinct excipulum not producing any crystals can be found in Farkas & Vězda 

(1993: 324). 

Byssoloma kalbii is known from only one locality in Madeira, where it 

colonizes the fronds of the fern Trichomanes speciosum, growing on the ground; 

other foliicolous species present on the same phorophyte are Byssoloma 

leucoblepharum and Porina leptosperma. The Riba do Seixal shelters the best


stand of laurisilva studied so far on Madeira; it is located at the lowest 

elevation seen for remnants of the laurisilva on the island. 

This new species is dedicated to Dr K. Kalb from Neumarkt (Germany) 

who, together with Prof. J. Hafellner, recently made an important contribution 

to the lichen flora of Madeira (Kalb & Hafellner 1992), including the 

description of several new taxa. 

Fellhanera seroexspectata Sérus. sp. nov. 

Foliicola Fellhanera species insignis thallo cum citrinis sorediis. 

Type: Portugal, Madeira, Casa de Queimadas, alt. 850–900 m, track towards Caldeirâo Verde, 

degraded laurisilva, on leaves of Lauraceae, mainly Laurus azorica and Persea indica, v 1992, 

Sérusiaux s.n. (LG—holotypus). 

(Fig. 4) 

Thallus epiphyllous, in the best preserved material formed of very tiny 

flattened greenish granules, tightly pressed against each other and forming a 

rugose surface when seen under high magnification (40), more loosely 

arranged at the thallus margin where they are separated by a photobiont-free 

whitish prothallus, usually dark green but sometimes greyish to pale brown, 

sometimes almost absent or reduced to a thin uniform film. Photobiont most 

probably a species of Chlorococcaceae, with spherical, green cells, 6–12 μm 

diam. Soralia always present, circular, 0·2–0·3(–0·4) mm diam., usually 

distributed all over the thallus, but sometimes almost confluent in its centre or 

irregularly arranged at its margins, rarely dissolving into a soredial mass that 

spreads over the thallus surface, flattened or slightly convex, in the best 

preserved material distinctly crateriform with a thin membrane maintaining 

the soredia together, citrine to greenish yellow when fresh, becoming more 

greenish when dry and almost losing any colour in old herbarium specimens 

(then dirty white). Soredia farinose, formed of several (2–10) glomerules, 

each containing one algal cell when young (10–15 μm diam.) and up to 10 

when older (15–20 μm diam.) surrounded by globose or cylindrical hyphal 

filaments, with a few short hyphal projections, without any pigmentation or 

crystals. 

Apothecia very rare, circular, 0·2–0·3(–0·5) mm diam., 0·15 mm in 

height, flat or slightly convex, with a dark brown disc, with a reddish hue and 

a non-prominent but persistent, paler margin. Excipulum well-developed, 

typically paraplectenchymatous, hyaline in sections, 50–60 μm thick under the 

hypothecium and less than 30 μm in lateral parts. Hypothecium brownish, 

slightly reddish, without any reaction in K. Hamathecium of coherent, simple 

or rarely branched paraphyses, slightly swollen at the spices, some of them 

containing the pigment present in the hypothecium. Asci of the Byssoloma-type 

(Hafellner 1984: 315), 50–6015 μm, 4–8-spored. Ascospores ellipsoid or 

nearly cylindrical, with rounded ends, 3–(4)-septate, not or very slightly 

constricted at the septa (even in a 10% KOH solution), with a thin gelatinous 

halo (best seen in KOH solution), (16–)19–225–7 μm. 

Pycnidia very rare, sessile, globose, slightly constricted at their base, 0·1– 

0·2 mm diam., wall dark bluish brown. Conidia usually numerous, bacilliform 

to ellipsoid but often deformed, sometimes slightly curved, 5–81–1·5 μm.


FIG. 4.Fellhanera seroexpectata (holotype). A, Thallus with apothecia and soredia; B, Sterile 

thallus; C, Thallus with pycnidia (arrows) and soredia. Scale=1 mm.


This new species clearly belongs to Fellhanera and is easily distinguished by 

the presence of soredia, a rare feature in foliicolous lichens. Although 

uncommon, it is easily detected in the field or in fresh material by its citrine or 

greenish yellow discrete soralia. The pigment that gives this characteristic 

colour has not been identified but the soralia do not react with K, C or P. The 

soredia can be described as consoredia as understood by Tønsberg (1992: 

34–37) as each of them consists of aggregated glomerules made of algal cells 

surrounded by globose to cylindrical hyphae. Although it could apply for each 

glomerule, the term goniocyst is avoided as it has been used for many different 

structures in the literature and I have suggested it should be restricted to the 

diaspores produced in some foliicolous Opegrapha species (Sérusiaux 1985). 

A comprehensive key to all described foliicolous species of Fellhanera is 

provided below. 

Fellhanera seroexspectata is known only from Madeira where it is reported 

from several laurisilva localities on the island. It does not require undisturbed 

forests as I collected it at Ribeiro Frio in a severely degraded spot near the 

tourist compound. It is not common but is usually seen as sterile thalli with 

scattered yellowish soredia. I first found this species in 1978 when examining 

a small collection of foliicolous lichens gathered at Ribeiro Frio by J. 

Duvigneaud, a Belgian botanist; the specimen has only three apothecia and did 

not provide enough material for a good type specimen. I had to wait until May 

1992 to find a suitable specimen; from my point of view, the epithet chosen to 

describe this attractive species (seroexspectata) is therefore appropriate. 

Additional specimens examined: Madeira: Ribeiro Frio, alt. 850 m, on Ruscus streptophyllus, vii 

1975, J. Duvigneaud 75.M.696 (LG); ibid., on leaves of Laurus azorica, xii 1979, Arvidsson (GB); 

ibid., ii 1988 and v 1992, Sérusiaux s.n. (LG, 3 specimens); Châo de Louros, N of Encumeada 

pass, alt. 800 m, ii 1988 and v 1992, Sérusiaux s.n. (LG, 3 specimens); Riba da Seixal, alt. 

300–400 m, v 1992, Sérusiaux s.n. (LG). 

Key to the Foliicolous Species of Fellhanera 

This key deals only with foliicolous species of Fellhanera Vězda, a typical 

representative of the Pilocarpaceae. The main apomorphy of this family, best 

represented in tropical and subtropical areas, is its ascus type, known as the 

Byssoloma-type (see Hafellner 1984: 315). 

Within the family, Fellhanera is characterized by the paraplectenchymatous 

structure of its excipulum: solely species with such an excipulum are accepted 

in the genus and therefore included in the key. The only exception is 

F. stanhopeae, the excipulum structure of which is impossible to distinguish as 

it is filled with crystals. Both species of Fellhaneropsis are, however, included 

for the absence of pycnidia may entail some confusion. Indeed the key is based 

on easily observable characters. 

The relationships between Fellhanera and Badimia Vězda, a genus suspected 

to belong to the Pilocarpaceae (Sérusiaux 1986: 6), are studied in detail by 

Lücking et al. (1994). Species that were mentioned as ‘ad int.’ by Lücking 

(1992: 136–138) and not formally described are not included in the key. 

I expect that many more species, either foliicolous or corticolous, will be 

discovered in the future, especially in tropical regions.


1 Ascospores 1-septate . . . . . . . . . . . . . . . . . . . . . . 2 

Ascospores with at least 3 septa . . . . . . . . . . . . . . . . . 7 

2(1) Pycnidia blue-grey, with a long beak, 0·15–0·3 mm long; apothecia 

0·15–0·5 mm diam., with a beige-brown to chocolate-brown disc 

and a persistent, beige, hardly prominent margin; ascospores 

9–12(–13)4–5 μm. Africa (Zimbabwe) . . . . . . . . . . . . 

Fellhanera vandenberghenii (Sérus.) Vězda 

Pycnidia, if present, without a long beak . . . . . . . . . . . . 3 

3(2) Apothecia black or brownish black, with a hardly distinct margin, 

0·2–0·25 mm diam.; epithecium with dark granules; ascospores 

10–123·5–4·5 μm. Africa (Zaïre) . . . . . . . . . . . . . . . 

Fellhanera encephalarti (Vězda) Vězda 

Apothecia not black, or, if dark brown, without any dark granules in 

the epithecium, with or without a distinct margin . . . . . . . 4 

4(3) Ascospores not exceeding 122·5 μm . . . . . . . . . . . . . 5 

Ascospores when mature always exceeding 122·5 μm . . . . . 6 

5(4) Apothecia very small, 0·08–0·2 mm diam., whitish to pale yellow, 

usually without any margin; hypothecium hyaline; ascospores 

6–112·5 μm. South America (Brazil) and Africa (Guinea) . . . 

Fellhanera parvula (Vězda) Vězda 

Apothecia larger, 0·2–0·4 mm diam., brownish red to dark brown, 

with a pale yellowish brown margin; hypothecium dark brown; 

ascospores 6–101·5–2 μm. Africa (Tanzania) . . . . . . . . . 

Fellhanera congesta (Vězda) Vězda 

6(4) Thallus usually distinctly farinose, continuous, always with a bluish 

tinge; apothecia 0·1–0·3 mm diam., with an orange disc, sometimes 

yellowish or reddish, and with a distinct, whitish, slightly 

prominent margin. Subcosmopolitan and locally abundant, 

ubiquitous (not strictly foliicolous) . . . . . . . . . . . . . . . 

Fellhanera bouteillei (Desm.) Vězda 

Thallus smooth or farinose, without any bluish tinge; apothecia 

0·1–0·2 mm diam., with a pale brown or lightly reddish brown 

disc, and a faintly visible whitish margin. Pantropical and rare . . 

Fellhanera semecarpi (Vainio) Vězda 

7(1) Ascospores 3-septate, very rarely 5-septate . . . . . . . . . . . 8 

Ascospores, when mature, with at least 5 septa, or variable but always 

with most ascospores with more than 3 septa, or submuriform 26 

8(7) Apothecia very small, 0·1–0·2 mm diam., convex and without any 

margin when mature, dark brown to almost black . . . . . . . 9 

Apothecia larger, very rarely less than 0·2 mm diam., convex or not 

when mature, margin present or not; apothecia colour dark brown 

or not . . . . . . . . . . . . . . . . . . . . . . . . . . . 10


9(8) Ascospores 12–163–5·5 μm; apothecia strongly convex when 

mature. Pantropical . . . . . . . . . . . . . . . . . . . . . . 

Fellhanera rhapidophylli (Rhem) Vězda 

=Bacidia myriocarpa Vězda 

Ascospores 18–25(–28)3–4 μm; apothecia slightly convex when 

mature. Western and central Europe, Tenerife and Madeira, a 

rather ubiquitous species, usually abundant on twigs . . . . . . 

Fellhaneropsis myrtillicola (Erichsen) Sérus. & Coppins 

10(8) Thallus formed of isolated, rarely confluent, whitish, and convex 

verrucae, or with a continuous thallus with distinct verrucae that 

may eventually break up into a soredial mass, or with a continuous 

thallus and typical soredia . . . . . . . . . . . . . . . . . 11 

Thallus never with typical soredia, nor with verrucae . . . . . 16 

11(10) Thallus formed only of isolated, rarely confluent, whitish and convex 

verrucae; apothecia growing on the thin pellucid prothallus or on 

the margin of the verrucae, 0·3–0·35 mm diam. with pale or 

dark brown disc and a thin but distinct paler margin. Australia 

(Queensland), Papua New Guinea and New Caledonia. [See 

note 1.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

Fellhanera bullata Kalb & Vězda 

Thallus not formed only of verrucae, but verrucae present or not over 

the thallus surface . . . . . . . . . . . . . . . . . . . . . 12 

12(11) Thallus continuous with whitish hemispherical verrucae, pale bluish 

grey; apothecia 0·25–0·4 mm diam. with a yellowish red disc and a 

white, sharply delimited margin. Central America (Costa Rica) . 

Fellhanera badimioides Lücking, Lumbsch & Elix 

Thallus with distinct verrucae that eventually break up into a soredial 

mass and then thallus greenish and apothecial disc dark brown, or 

with soredia . . . . . . . . . . . . . . . . . . . . . . . . 13 

13(12) Thallus with distinct verrucae that eventually break up into a soredial 

mass, rarely with large and sometimes confluent soralia that spread 

over the thallus surface . . . . . . . . . . . . . . . . . . . 14 

Thallus never with verrucae, but with soredia . . . . . . . . . 15 

14(13) Excipulum encrusted with colourless crystals (the paraplectenchymatous 

structure of the excipulum is therefore indistinct); thallus 

with verrucae of various size, that frequently break up into a coarse 

soredial mass; thallus smooth to farinose. Distribution: see note 2 

Fellhanera stanhopeae (Müll. Arg.) Lücking, Lumbsch & 

Elix 

Excipulum not encrusted with crystals, with a distinct paraplectenchymatous 

structure; thallus with verrucae that very frequently 

break up into a soredial mass, or with abundant, large, yellowish 

soralia that are often confluent and cover large parts of the thallus;


thallus farinose to coarsely farinose. Africa (Zaïre and Rwanda). 

[See note 3] . . . . . . . . . . . . . . . . . . . . . . . . . . 

Fellhanera lambinonii (Sérus.) Lücking & Sérus. 

=Bacidia michaeliana Sérus. 

15(13) Thallus smooth, continuous, whitish or brownish, reacting K+ red 

brown, with a distinct dark violaceous prothallus; soredia whitish, 

punctiform, loose and dispersed, becoming confluent and larger on 

older thalli; apothecia 0·4–0·7 mm diam., with a pale orange to 

yellowish brown disc and a pale orange distinct margin that 

eventually vanishes; ascospores 13–164–4·5 μm. Africa (Zaïre 

and Ivory Coast) . . Fellhanera sorediantha (Vězda) Vězda 

Thallus rugose under high magnification and formed of packed 

greenish granules in well-preserved material, sometimes absent or 

formed by a thin brownish membrane; soredia citrine to greenish 

yellow, losing their colour in herbaria, rarely confluent and 

coalescing; apothecia very rare, 0·2–0·3(–0·5) mm diam., with a 

dark brown disc and a paler, distinct and persistent margin; 

ascospores (16–)19–225–7 μm. Macaronesia (Madeira) . . . . 

Fellhanera seroexspectata Sérus. 

16(10) Apothecia almost black, or brown, possibly with a red tinge but never 

ochre, nor with a vivid red colour. . . . . . . . . . . . . . 17 

Apothecia paler, never blackish or brown . . . . . . . . . . . 22 

17(16) Ascospores not exceeding 13 μm in length and 2·5 μm in width 

(8–132–2·5 μm); apothecia 0·3–0·8 mm diam. with a red-brown 

disc and a pale reddish margin, at first distinct and hardly 

prominent but soon disappearing; hypothecium dark-brown, K+ 

purple-red. Africa (Zaïre). Fellhanera wirthii (Vězda) Vězda 

Ascospores always exceeding 13 μm in length and 2·5 μm in width 

when mature; hypothecium K+ purple-red or not. . . . . . 18 

18(17) Hypothecium brownish red, usually K+ purple-red, rarely K+ 

brown; apothecia 0·3–0·5 mm diam. with a brownish red disc, at 

first with a distinct, slightly prominent, pale reddish brown margin; 

ascospores 18–253·5–4 μm. Australia (Queensland and New 

South Wales) Fellhanera endopurpurea Hafellner & Vězda 

Hypothecium never reacting K+ purple-red, even when dark brown 

or brownish . . . . . . . . . . . . . . . . . . . . . . . . 19 

19(18) Apothecia mainly developed on a hypophyllous, marginal, nonlichenized 

mycelium, 0·3–0·5 mm diam.; disc first plane, soon 

convex, bluish grey to blackish brown, usually with a greyish white 

pruina, with a pale greyish margin, at first prominent but soon 

thinner and almost disappearing in old apothecia; ascospores 

17–273·5–5 μm. Central America (Cocos Is) . . . . . . . . . 

Fellhanera avilezii Lücking 

Apothecia mainly epiphyllous, developed on lichenized thallus . 20


20(19) Thallus green to pale green; apothecia 0·3–0·45(–0·5) mm diam., 

with a dark brown to carbonaceous black disc and a concolorous or 

slightly paler margin which at first is thick and prominent but 

becomes thinner and not prominent; pruina on the disc absent 

or, if present, very thin and bluish; ascospores 14–173–5 μm. 

Pycnidia usually present, black. Europe (Italy/Calabria) and 

Macaronesia (Madeira and Tenerife) . . . . . . . . . . . . . . 

Fellhanera christiansenii Sérus. & Vězda 

Thallus greenish grey or bluish white; apothecia and pycnidia 

different . . . . . . . . . . . . . . . . . . . . . . . . . . 21 

21(20) Thallus greenish grey; apothecia 0·2–0·4 mm diam., with a grey to 

livid black disc and a distinct, whitish, hardly prominent margin; 

pruina on the disc whitish, always present on young apothecia; 

ascospores 15–203·5–4 μm. Pycnidia unknown. Central 

America (Costa Rica, Guatemala, St. Lucia) . . . . . . . . . . 

Fellhanera santessonii Barillas & Lücking 

Thallus bluish white, or rarely bluish grey; apothecia 0·2–0·4 mm 

diam., with a dark brown disc and a distinct but thin, nonprominent, 

pale brown margin; pruina on the disc very rare; 

ascospores 12–183·5–5·5 μm. Pycnidia rare, pale yellowish grey. 

Pantropical but mostly common in Africa . . . . . . . . . . . . 

Fellhanera sublecanorina (Nyl.) Vězda 

22(16) Hypothecium K+ red-brown or red. . . . . . . . . . . . . . 23 

Hypothecium K . . . . . . . . . . . . . . . . . . . . . . 24 

23(22) Apothecia 0·4–0·5(–0·6) mm diam., with an orange-red to brownish 

disc and a pale orange, thin, non-prominent margin; thin whitish 

pruina sometimes present on the disc; excipulum sometimes 

filled with small yellowish brown granules that dissolve in K; 

ascospores 15–204·5–5 μm. Central America (Costa Rica and 

Guadeloupe) and Africa (Zaïre) . . . . . . . . . . . . . . . . 

Fellhanera lisowskii (Vězda) Vězda 

Apothecia 0·3–0·4 mm diam., with an orange-red disc and a distinct, 

pale orange or white, at first thick but soon non-prominent margin; 

excipulum and hypothecium hyaline but K+ vivid yellow to red; 

ascospores 12–154–4·5 μm. North America (USA—Florida) 

and Africa (Guinea and Zaïre) . . . . . . . . . . . . . . . . . 

Fellhanera aurantiaca (Vězda) Vězda 

24(22) Thallus granular, grey green to ochraceous; apothecia 0·2–0·4 mm 

diam., with a pale orange, sometimes pruinose disc and a thin, 

non-prominent, very pale orange to almost white margin; ascospores 

11–162·5–4·5 μm; apical parts of paraphyses with 1–3 

irregular swellings; pycnidia frequent. NW Europe, mostly corticolous, 

but sometimes growing on Vaccinium or Buxus leaves, or 

on Picea or Abies needles . . . . . . . . . . . . . . . . . . . . 

Fellhanera subtilis (Vězda) Diederiech & Sérus.


Thallus farinose, very thin, bluish grey to bluish white; apical parts of 

paraphyses without swellings . . . . . . . . . . . . . . . . 25 

25(24) Apothecial margin at first thick, prominent, pale orange and very 

minutely pruinose, then becoming thinner and almost nonpruinose; 

paraphyses sinuous, branched and anastomosed and 

2 μm thick; apothecia 0·2–0·4 mm diam., with an orange to 

pinkish orange disc; hypothecium brownish; ascospores 

14–194–5 μm. Central America (Trinidad) and Africa 

(Guinea) . . . . . . . . . Fellhanera carnea (Vězda) Vězda 

Apothecial margin thin, non-prominent, almost white and epruinose; 

paraphyses almost simple, straight and 1 μm thick; apothecia 

0·2–0·4 mm diam., with a pinkish orange to pale brown disc; 

hypothecium pale brownish; ascospores 10–152·5–4·5 μm. 

Pantropical but rare . . Fellhanera subternella (Nyl.) Vězda 

26(7) Ascospores 2–8 per ascus; mature ascospores of various shapes and 

irregularly with (3–)5–7(–9) transverse septa and sometimes with a 

longitudinal septum, 15–374·5–8 μm; apothecia 0·3–0·4 mm 

diam., with a red brown disc and a thin, orange-red, slightly 

prominent margin. Central America (Costa Rica) and Africa 

(Guinea and Tanzania). Fellhanera paradoxa (Vězda) Vězda 

Ascospores mostly 8 per ascus, of regular shape . . . . . . . . 27 

27(26) Ascospores submuriform, with 7–9 transverse and 1–3 longitudinal 

septa, 26–335–10 μm; apothecia 0·3–0·5 mm diam., with a 

yellowish or reddish brown disc and a thin, pale brown, slightly 

prominent margin. Central America (Costa Rica and St. Vincent) 

and South America (Brazil). . . . . . . . . . . . . . . . . . . 

Fellhanera elliottii (Vain.) Vězda 

Ascospores (3–)5–7-septate, never submuriform . . . . . . . . 28 

28(27) Ascospores (3–)4–7-septate, acicular-fusiform, usually with one end 

attenuated and more or less irregularly curved, 30–35(–42)3– 

4·5 μm; apothecia 0·2–0·4 mm diam., with a flat, livid brown to 

pinkish brown disc, at first with a thick, slightly raised margin, 

eventually immarginate, convex and tuberculate. Western and 

central Europe, and Macaronesia (Madeira); corticolous, occasionally 

found on leaves (in Madeira only) . . . . . . . . . . . 

Fellhaneropsis vezdae (Coppins & P. James) Sérus. & 

Coppins 

Ascospores mostly with a regular number of septa (5 or 7), ellipsoid 

to fusiform; sometimes with one end attenuated and slightly 

curved (F. microdiscus); apothecia never becoming tuberculate 29 

29(28) Ascospores 7-septate, 18–244–4·5 μm; apothecia 0·2–0·4 mm 

diam., with a brown disc and a thin, pale brown, slightly prominent 

margin. Central and South America (where the species is mostly


abundant), Australia (New South Wales) and Africa (Ivory Coast 

and ? Tanzania [see note 4]) . . . . . . . . . . . . . . . . . . 

Fellhanera dominicana (Vainio) Vězda 

Ascospores 5-septate, rarely a few ascospores with 6 or 7 septa 30 

30(29) Most ascospores exceeding 25 μm in length [24–32(–34)4–5 μm]; 

pruina present, at least in young apothecia . . . . . . . . . 31 

Most ascospores not reaching 25 μm in length [14–263–5 μm]; 

pruina never present, even in young apothecia . . . . . . . 32 

31(30) Apothecia developed on an epiphyllous, lichenized thallus; apothecia 

0·2–0·4 mm diam., with a brownish disc, sometimes almost black 

or with a distinct red tinge, when young usually with a thin whitish 

pruina, and with a distinct pale brown margin. Central America 

(Costa Rica and Cuba) . . . Fellhanera winkleriana Lücking 

Apothecia mostly developed on a hypophyllous, marginal, unlichenized 

mycelium; apothecia 0·5–0·6 mm diam., with a dark brown 

disc; whitish to bluish pruina present on the disc and on the margin 

in young apothecia, but thick and persistent only on the margin. 

Central America (Cuba) . . . . . . . . . . . . . . . . . . . . 

Fellhanera ekmanii (Vězda) Lücking 

32(30) Apothecial disc pale to dark brown; margin pale brown, thin and 

non-prominent; ascospores bacilliform to ellipsoid, usually not 

curved and not tapering towards one end. Pantropical but mostly 

abundant only in Africa . . . . . . . . . . . . . . . . . . . . 

Fellhanera fuscatula (Müll. Arg.) Vězda 

Apothecial disc red-brown; margin pale reddish brown, thin and 

non-prominent; ascospores ellipsoid, usually slightly curved and 

attenuated at one end. Asia (Philippines and Vietnam) and Australia 

(Queensland). Fellhanera microdiscus (Vainio) Vězda 

Notes 

1. Dr K. Kalb has recently collected Fellhanera bullata in New Caledonia: Prov. Sud, Mts 

Koghis-Dumbéa, 550 m alt., tropical rain forest, 23 viii 1994, K. & A. Kalb (hb. Kalb!). 

A collection from that country has just been distributed in Lücking Lich. Fol. Exs. No. 136 

(1995). The species is also present in Papua New Guinea: Eastern Highlands prov., 

Gahavisuka Provincial Park, 2300–2450 m, mossy mountain forest, 11 viii 1992, Sérusiaux 

13762-45 (LG). 

2. Fellhanera stanhopeae is said to be present in Central and South America and in Africa by 

Santesson (1952: 474–475) and by Vězda (1980: 93) but typical specimens, examined by R. 

Lücking and myself, are all from the Neotropics. Those from Africa may all belong to 

F. lambinonii. Bacidia stanhopeae has been transferred to Badimia by Vězda (in Lich. Sel. Exs. 

No. 2307, 1989) but does not belong there. 

3. Fellhanera lambinonii (Sérus.) Lücking & Sérus. comb. nov. Bas.: Bacidia lambinonii Sérus., 

Lejeunia N.S., 90: 12 (1978). 

Bacidia michaeliana Sérus. represents the sorediate form of that species. Several collections 

(LG) that were not available in 1978 for the original description of this taxon (Sérusiaux 

1978: 15–16) show all intermediate stages between typical Bacidia lambinonii and Bacidia 

michaeliana. 

4. Fellhanera dominicana is mentioned as ‘cf. dominicana’ from Tanzania by Vězda (1975: 417).


Preliminary List of Foliicolous Lichens and Their Lichenicolous 

Fungi from Madeira 

Ampullifera foliicola Deighton 

This lichenicolous hyphomycete was first mentioned from the island by 

Arvidsson & Wall (1985: 42) and by M. S. Christiansen (in Santesson 1986: 

1) on the thallus of Byssoloma subdiscordans. It is indeed common on this 

abundant foliicolous species and was found only on it. When abundant it can 

hamper the development or production of its host’s apothecia and spreads 

over the leaf surface, but it is nevertheless unable to colonize other lichen 

species. It is also present on the same species in the French Pyrenees. 

Ampullifera foliicola is probably a pantropical species, and interestingly grows 

over several different hosts in other parts of the world. 

Arthonia muscigena Th.Fr. 

This species is thoroughly described in Purvis et al. (1992: 84) and is usually 

known as A. leucodontis (Poelt & Döbbeler) Coppins, which is a synonym. It 

is a rather common, ubiquitous species of lowland western Europe and is 

tolerant of suburban conditions. It is not surprising to find it on leaves. 

Foliicolous specimens have, however, been named A. microsticta Vain. (Vězda 

Lich. Sel. Exs. No. 1676, 1980). The type collection of this forgotten species 

(West Indies, Dominica, Elliott 517, BM!) is a small leaf fragment with several 

apothecia of the Arthonia growing over the leaf surface, or over a smooth 

bluish lichen thallus. This group of tiny Arthonia with 1-septate ascospores is 

very little known and it cannot be ruled out that A. microsticta is just a form of 

the widespread A. muscigena Th.Fr. More material from the West Indies is 

needed to understand correctly this taxon. 

In Madeira, Arthonia muscigena grows directly on the leaf surface or forms 

its own very thin greenish thallus; it is locally abundant. In western Europe 

(S Belgium, S France, Spanish side of the Pyrenees), it specializes on the 

goniocysts of Woessia species on Buxus leaves and twigs; it does not seem to 

damage its host. 

Aulaxina sp. 

A sterile specimen collected in 1988 at Châo do Louros with numerous 

hyphophores definitely belongs to that genus; the hyphophores are typical of 

the pantropical Aulaxina quadrangula (Stirt.) R. Sant. but the absence of 

ascomata precludes any definite identification. 

Byssoloma aptrootii Sérus. 

This recently described species (Sérusiaux 1993: 451–454) is one of the 

most common foliicolous lichen in the laurisilva of Madeira, at least in the less 

disturbed localities. Outside Madeira, it is known only from Tenerife and 

Gomera where it is also common. 

Byssoloma leucoblepharum (Nyl.) Vain 

A common species in Madeira, also found on twigs and bark. It is a 

common foliicolous lichen in SW Europe on Buxus leaves and is also found on


bark, especially on Corylus and on Quercus. Its general distribution can be 

described as pantropical, expanding into warm temperate regions. 

Byssoloma marginatum (Arnold) Sérus. 

Tapellaria similis Kalb in Kalb & Hafellner, Herzogia 9: 90 (1992). 

This species is known mainly as corticolous in western and central Europe, 

and as foliicolous and corticolous in Madeira and in Tenerife. So far I have 

seen specimens from Germany (Bavaria), Austria (Steiermark), the British 

Isles, S and SW France, Spain/Galicia (see Lopez de Silanes & Carballal 1991: 

48; specimen seen by myself), from Portugal (van den Boom et al. 1990: 470; 

the specimen mentioned as ‘Byssoloma sp.’ from the Serra de Sintra belongs to 

B. marginatum), from Italy/Calabria (the species is present on living needles 

and on bark of Abies in a set of lichens sent to me by D. Puntillo, but it is not 

mentioned in Puntillo & Vězda 1994) and from Tenerife (where it is common 

on Erica in the ‘Fayal-Brezal’, together with Byssoloma leucoblepharum and 

Micarea pycnidiophora Coppins & James). The species is also mentioned from 

S Sweden by Santesson (1993: 42). In Madeira, it is not uncommon on leaves 

but never develops conspicuous thalli; it also grows on twigs and on bark of 

several trees in the laurisilva. Tapellaria similis Kalb, just described on bark of 

Laurus azorica in Madeira (Kalb & Hafellner 1992: 90), is typical Byssoloma 

marginatum (holotype: K. Kalb 23296, hb. Kalb!). 

Byssoloma subdiscordans (Nyl.) James 

This is one of the most common foliicolous lichens in Madeira, also 

colonizing the living leaves of introduced species in suitable conditions, and 

rarely epiphytic on twigs or on bark. It was reported from Madeira by 

Santesson (1952: 493) and is a widely distributed species in the tropics as well 

as in temperate Europe where it is known as either saxicolous, corticolous on 

spruce and fir twigs, or foliicolous on spruce and fir needles and on Buxus 

leaves. 

Dimerella luteola Kalb 

This recently described species (Kalb & Hafellner 1992: 62–63) is the 

Madeiran vicariant of the European D. lutea (Dickson) Trevisan. It is a 

common epiphytic species in the laurisilva and is occasionally present on living 

leaves, especially at Seixal. 

Fellhanera bouteillei (Desm.) Vězda 

This is one of the most widespread species on the planet, being present in 

boreal, temperate and tropical areas, growing on coastal rocks, on twigs, on 

bark and on living leaves. It was known in Madeira by Santesson (1952: 434) 

where it is abundant on leaves. 

Fellhanera christiansenii Sérus. & Vězda 

The abundance of this species in Madeira is comparable with that of 

Byssoloma aptrootii with which it often grows. I first collected it in 1988 in large


quantities and was ready to describe it as new (under the name Fellhanera 

nigra) when Dr A. Vězda sent to me a preliminary version of a manuscript 

dealing with new taxa of foliicolous lichens, including this new species of 

Fellhanera. It was then decided to describe it together under the epithet 

christiansenii. I have, however, spread the epithet nigra for several years and the 

curators of herbaria should be aware that ‘F. nigra Sérus. ad int.’ is the same 

as F. christiansenii Sérus. & Vězda (in Vězda 1994: 130–131). The species is 

described on a collection from Tenerife where it is common; it is also known 

from Calabria in Italy (Puntillo & Vězda 1994; several specimens checked by 

myself). The reports of Fellhanera buxi (Vězda & Vivant) Vězda from Tenerife 

by Lumbsch & Vězda (1992: 25) are all based on typical specimens of 

F. christiansenii (both specimens cited seen). 

Gyalectidium colchicum Vězda 

This species was described by Vězda (1983: 58–60) from collections made 

in the western Caucasus (Georgia and Russia). The foliicolous populations of 

Gyalectidium from Madeira are referred to that species as they have hyphophores 

typical of the genus (see Sérusiaux & De Sloover 1986: 282–289) and 

identical with those of that species [size: 0·15–0·2(–0·3) mm in total length]. 

They do not belong to G. filicinum Müll. Arg. as claimed by Arvidsson & Wall 

(1985: 42; specimens mentioned by these authors examined, GB!). Vězda 

(1983: 58–60) has described pycnidia in that species; they are also present in 

Madeira. They are seen as slightly raised, dark bluish spots on the thallus 

surface, and their conidia are bacilliform to slightly bifusiform and measure 

2–30·75 μm. Apothecia are absent in the Caucasus material but are 

sometimes present in Madeira; they are typical of the genus and are seen as 

rounded sessile discs, 1–2(–10) per thallus with a pale grey or green disc, 

0·2–0·3 mm diam., and the ascospores are single in the asci, muriform and 

measure 32–3813–18 μm. The species is sometimes attacked by the 

hyphomycete Hansfordiellopsis lichenicola (see under that species). Gyalectidium 

colchicum is also present in the Azores but is unknown in the Canary Islands. 

It is the only species of the genus present in Madeira, whereas the recently 

described G. setiferum Vězda & Sérus. (in Sérusiaux 1993: 454–458) and 

G. caucasicum (Elenkin & Woron.) Vězda are observed in continental Europe, 

the latter being extremely rare in Western Europe (two localities known: 

France/Atlantic Pyrenees and Spain/Cataluñya). 

Hansfordiellopsis lichenicola (Batista & Maia) Deighton 

This lichenicolous hyphomycete is widespread in tropical areas on the three 

continents (see Hawksworth 1979: 227), infecting several species, mostly 

those with setae. In Madeira it is strictly confined to Gyalectidium colchicum 

and when abundant is able to damage its host; it is also present on the same 

species in the western Caucasus. 

Porina hoehneliana (Jaap) R. Sant. 

This species is very common in western Europe [France, Spain (Navarra 

and Cataluñya), Croatia], and in the western Caucasus (Russia and Georgia)


and has also been reported in the Himalaya range (Vězda & Poelt 1988: 425). 

In Madeira I have found it in one locality only, the Seixal laurisilva, where it 

is very common. This locality shelters the only laurisilva at low elevation 

(300–400 m vs 800–850 m for the other localities). It was, however, found at 

Châo de Louros by C. Tavares in 1951 and published by Santesson (1952: 

260) under Porina semecarpi Vainio, which is absent from the island. 

Porina leptosperma Müll. Arg. 

The species was already mentioned in Madeira by Santesson (1952: 258); 

it is common on the island. It is also abundant in the western Pyrenees in 

France, where it has been confused with P. hoehneliana; the No. 1083 and 

1084 of the Vězda Lich. Sel. Exs., collected in France on Buxus leaves and 

on Ruscus cladodes and distributed as P. hoehneliana, represent typical 

P. leptosperma. The species is otherwise pantropical. 

Psoroglaena stigonemoides (Orange) Henssen 

This species is common in western Europe [especially as an epiphyte on 

Sambucus, frequently associated with Anisomeridium nyssaegenum (Ellis & 

Everh.) Harris] and has just been reported from Madeira (Kalb & Hafellner 

1992: 70). It is a common epiphytic and muscicolous species in the laurisilva 

of the island and is occasionally present on living leaves. The same situation 

occurs in SW France where the species is locally extremely abundant and can 

invade Buxus twigs and leaves. The taxonomic affinities of this species have 

just been established by Henssen (1995) and are accepted here. 

Strigula angustata Sérus. & Roux ad int. 

This species is close to Raciborskiella minor Vězda, described from Georgia 

in the western Caucasus (Vězda 1983: 49–51), and widespread on Buxus twigs 

in southern France. It is more closely related to the bulk of Strigula species 

than to the type species of Raciborskiella [R. janeirensis (Müll. Arg.) R. Sant.]. 

The status of S. angustata is currently being studied with Dr C. Roux (see 

Roux & Bricaud 1993) and this taxon may end as a variety of R. minor as the 

only diagnostic character is the size of macroconidia. It is known from 

Madeira and from two localities in France (Lot and Vercors). 

Strigula nitidula Mont. 

This species is reported from Madeira by Santesson (1952: 182) and is 

indeed common on the island. Specimens with an effigurate thallus and 

distinct lobes and a thin black line at the margins come close to S. concreta 

(Fée) R. Sant., a pantropical species. Following Lücking (1992: 39), they 

nevertheless belong to S. nitidula: S. concreta has a thickened thallus and never 

has a metallic thallus with scattered small black points. Strigula nitidula is 

common in SW Europe, reaching Brittany in W France (Josien 1967: 829, 

de Foucault et al. 1982: 75). 

Tapellaria epiphylla (Müll. Arg.) R. Sant. 

Santesson (1952: 506) reports this species in the Azores and in Madeira 

where it is one of the most common foliicolous species. It also grows on living


leaves of introduced species (e.g. Camellia) in suitable localities. In the Canary 

Islands, it is known only in Gomera. Tapellaria epiphylla is a common 

foliicolous species in the Neotropics and in Africa. 

Vezdaea dawsoniae Döbbeler 

This very tiny pedicellate Vezdaea was first described from Papua New 

Guinea (where it is common in the mountains, pers. obs.) and was later 

reported from Cuba (Vězda 1984: 192; as ‘cf. dawsoniae’), from the Pyrenees 

in France (Sérusiaux 1989: 90) and from the Caucasus (Giralt et al. 1993: 

mentioned in the key page 716, but not in the conspectus on page 720; for 

unknown reasons, Vezdaea obscura, described by Döbbeler 1981: 461–464 

from Tasmania and New Guinea, is not studied in that survey of the genus). 

In Madeira, V. dawsoniae has been found as foliicolous (in small quantities) 

only at Châo do Louros. 

Woessia apiahica (Müll. Arg.) Sérus. comb. nov. 

Patellaria apiahica Müll. Arg., Lichenes epiphylli novi: 9, 1890. 

Bacidia apiahica (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 4: 174, 1926. 

Bacidina apiahica (Müll. Arg.) Vězda, Folia Geobot. Phytotax. (Praha) 25: 432, 1990. 

The generic name Woessia Hawksw. & Poelt antedates Bacidina Vězda and 

is now used for the rather well-delimited group of species of Bacidia s.l. with 

a thallus made of goniocysts s.lat., pale orange to dark brown apothecia, a 

typically paraplectenchymatous excipulum, simple paraphyses with swollen 

apices, acicular ascospores spirally arranged in the asci and filiform-sigmoid 

conidia forming a coiled mass in the pycnidia (Sérusiaux 1995). The necessary 

combinations of the species mentioned in this paper are introduced. Woessia 

apiahica was already mentioned in Madeira by Santesson (1952: 443) and 

indeed Woessia with small pale orange apothecia are common on leaves in 

the island. The status of those populations as well as the genuine identity of 

W. apiahica are, however, still a great challenge to me. In Madeira I suspect 

that two taxa are involved and I am quite sure that Woessia vasakii (Vězda) 

Sérus. comb. nov. [Bacidia vasakii Vězda, Folia Geobot. Phytotax. (Praha) 

18: 64, 1983], which is abundant in southern France, is not present. 

Woessia canariensis (Lumbsch & Vězda) Sérus. comb. nov. 

Bacidina canariensis Lumbsch & Vězda, Lichenologist 24: 22, 1992. 

I first collected this species in Madeira in 1988 where it is rather rare; in 

contrast it is more frequent in Tenerife where I gathered large samples, 

including several with almost black apothecia. The species prefers rather dry 

laurisilva, a niche which is more frequent in Tenerife than in Madeira. The 

type collection (hb. Lumbsch!) comes from the Anaga Mts in Tenerife 

(Lumbsch & Vězda 1992: 22–24). One of the specimens mentioned by 

Lumbsch & Vězda (Tenerife, Las Montanas de Anaga, 25 iii 1976, Santesson 

26837, hb. Lumbsch!) contains only Byssoloma aptrootii.


Species to be Excluded from the Madeiran Flora 

Dimerella epiphylla (Müll. Arg.) Malme 

This common pantropical species is reported from Madeira by Follmann 

(1990: 100). The only specimen cited (Ribeiro Frio, vi 1987, S. Schuhmacher, 

KOELN 33244) has not been examined, but another one, also collected at 

Ribeiro Frio in June 1987 by S. Schuhmacher and annotated Dimerella 

epiphylla (No. 708, KOELN), has been studied: it is a pale but nevertheless 

typical Fellhanera bouteillei. Dimerella epiphylla has not been found in my own 

collections. It should probably be removed from the Madeiran flora. 

Dimerella lutea (Dicks.) Trevisan 

This species is reported as foliicolous in Madeira by Santesson (1952: 403) 

on the basis of a collection made by Tavares. R. Santesson had a rather wide 

concept of that species, including the now easily distinguished Dimerella 

fallaciosa (Müll. Arg.) Vězda. The specimen is preserved at LISU and has been 

examined (Châo do Louros, 31 vii 1951, C. Tavares No. 4428-Q): it contains 

only two apothecia. Therefore I have not made any section of them; I suspect 

it is a foliicolous Dimerella luteola Kalb, which is a common epiphytic species 

in the laurisilva of the island and which is occasionally present on living leaves. 

Porina semecarpi Vain. 

The species is reported from Madeira by Santesson (1952: 260); the single 

mentioned collection is preserved at LISU and has been examined (Entre a 

Encumeada e a Châo do Louros, 31 vii 1951, C. Tavares No. 4428-F): it is a 

typical Porina hoehneliana. Porina semecarpi must therefore be removed from 

the Madeiran flora. 

Tricharia triseptata R. Sant. 

The only specimen cited by Follmann (1990: 101) for this species in 

Madeira has been studied (v 1987, S. Schuhmacher KOELN 33258); it 

contains only unlichenized fungi amongst which Dennisiella babingtonii (Berk.) 

Bat. & Cif. is abundant and has obviously been confused with Tricharia 

triseptata. This species should therefore be removed from the Madeiran flora. 

Dennisiella babingtonii is a member of the Coccodiniaceae, widespread worldwide 

on living leaves, especially in humid areas; it is very common in Madeira 

(description in Batista & Ciferri 1962: 37–38). 

I would like to thank the curators of the following institutions and the colleagues who keep private 

herbaria which allowed me to study material in their care: B, BM, E, GB, GZU, H, KOELN, LD, 

LISU, S, UPS, hb. A. Aptroot, hb. S. Christiansen, hb. P. Diederich, hb. J. Etayo, hb. T. 

Lumbsch, hb. K. Kalb, hb. D. Puntillo, hb. L. Spier and hb. A. Vězda. Dr C. Printzen kindly 

arranged the loan of type material of Lecidea quintula Nyl. Dr B. J. Coppins is warmly thanked for 

most valuable comments on the species treated in this paper and for reading and improving the 

manuscript. Drs U. Arup, P. Diederich, R. Lücking and Prof. J. Lambinon also read the 

manuscript with great care and made several interesting suggestions and remarks. 

REFERENCES 

Arup, U. & Ekman, S. (1994) Tre lavar i släket Fellhanera på blåbär. Svensk Botanisk Tidskrift 88: 

33–41.


Arvidsson, L. & Wall, S. (1985) Contribution to the lichen flora of Madeira. Lichenologist 17: 

39–49. 

Barillas, R. & Lücking, R. (1992) Líquenes foliícolos de Guatemala. Un estudio taxonómico 

preliminar. Cryptogamie, Bryologie Lichénologie 13: 297–317. 

Batista, A. C. & Ciferri, R. (1962) The Chaetothyriales. Beihefte zur Sydowia, Annales Mycologici, 

Ser. III: 1–129. 

Benfield, B. (1992) In New, rare, and interesting British lichens records. British Lichen Society 

Bulletin 71: 42. 

Boom, P. P. G. van den, Aptroot, A. & Knapp, W. O. van der (1990) New or interesting lichen 

records from Portugal. Nova Hedwigia 50: 463–472. 

Bricaud, O., Roux, C., Ménard, T. & Coste, C. (1993) Champignons lichénisés et lichénicoles de 

la France méridionale: espèces nouvelles et intéressants (8). Bulletin de la Société linnéenne de 

Provence 44: 99–110. 

Coppins, B. J. (1983) A taxonomic study of the lichen genus Micarea in Europe. Bulletin of the 

British Museum (Natural History), Botany series 11: 17–214. 

Coppins, B. J. & James, P. W. (1978) New or interesting British lichens II. Lichenologist 10: 

179–207. 

Diederich, P., Sérusiaux, E. & Boom, P. van den (1991) Lichens et champignons lichénicoles 

nouveaux ou intéressants pour la flore de Belgique et des régions voisines. V. Lejeunia N.S. 

136: 1–47. 

Döbbeler, P. (1981) Moosbewohnende Ascomyceten. V. Die auf Dawsonia vorkommenden 

Arten der Botanischen Staatssammlung München. Mitteilungen der Botanischen 

Staatssammlung München 17: 393–474. 

Farkas, E. & Vězda, A. (1993) Five new foliicolous lichen species. Folia Geobotanica Phytotaxonomica 

(Praha) 28: 321–330. 

Follmann, G. (1990) Zur Kenntnis der Flechtenflora und Flechtenvegetation von Madeira und 

den umliegenden Inseln. I. Chorologisch-soziologischer Abriss. Courier Forschungsinstitut 

Senckenberg 129: 91–102. 

de Foucault, B., Sérusiaux, E. & Van Haluwyn, C. (1982) Une nouvelle station française de 

lichens foliicoles dans le Massif central occidental (Aveyron). Cryptogamie, Bryologie 

Lichénologie 3: 73–76. 

Giralt, M., Poelt, J. & Suanjak, M. (1993) Die Flechtengattung Vezdaea mit V. cobria spec.nov. 

Herzogia 9: 715–724. 

Hafellner, J. (1984) Studien in Richtung einer natürlichen Gliederung der Sammelfamilien 

Lecanoraceae und Lecideaceae. Beihefte Nova Hedwigia 79: 241–371. 

Hafellner, J. (1992) A New Checklist of Lichenized and Lichenicolous Fungi of the Madeira 

Archipelago. Graz: Institut für Botanik der Karl-Franzens Universität. 

Hansen, A. & Sunding, P. (1993) Flora of Macaronesia. Checklist of vascular plants. 4th revised 

edition. Sommerfeltia 17: 1–295. 

Hawksworth, D. L. (1979) The lichenicolous Hyphomycetes. Bulletin of the British Museum 

(Natural History), Botany series 6: 183–300. 

Henssen, A. (1995) Psoroglaena costaricensis, a new lichen from Costa Rica and remarks on other 

taxa of the genus Psoroglaena (Verrucariaceae). Bibliotheca Lichenologica 57: 199–210. 

Jacobsen, P. (1992) Flechten in Schleswig-Holstein: Bestand, Gefährdung und Bedeutung als 

Bioindikatoren. Mitteilungen der Arbeitsgemeinschaft Geobotanik in Schleswig-Holstein und 

Hamburg 42: 1–234. 

Jacobsen, P. & Coppins, B. J. (1989) On the identity of some ‘endemic’ North German lichens. 

Nova Hedwigia 49: 255–273. 

Josien, M. (1967) (‘1966’) Strigula nitidula Mont., lichen épiphylle en France. Revue Bryologique 

et Lichénologique 34: 829–830. 

Kalb, K. & Hafellner, J. (1992) Bemerkenswerte Flechten und lichenicole Pilze von der Insel 

Madeira. Herzogia 9: 45–102. 

Kalb, K. & Vězda, A. (1990) Die Flechtengattung Byssoloma in der Neotropis (eine 

taxonomische-phytogeographische Studie). Nova Hedwigia 51: 435–451. 

Lopez de Silanes, M. E. & Carballal, R. (1991) Líquenes epífitos de la fraga de Caaveiro (La 

Coruña: Espana), II. Cryptogamie, Bryologie Lichénologie 12: 47–54. 

Lücking, R. (1992) Foliicolous lichens—a contribution to the knowledge of the lichen flora of 

Costa Rica, Central America. Beiheft Nova Hedwigia 104: 1–179.


Lücking, R., Lumbsch, H. T. & Elix, J. A. (1994) Chemistry, anatomy and morphology of 

foliicolous species of Fellhanera and Badimia (Lichenized Ascomycotina: Lecanorales). 

Botanica Acta 107: 393–401. 

Lumbsch, H. T. & Vězda, A. (1992) Contributions to the lichen flora of Tenerife. Lichenologist 24: 

21–26. 

Poelt, J. & Vězda, A. (1990) U} ber kurzlebige Flechten. Bibliotheca Lichenologica 38: 377–394. 

Poelt, J. & Vězda, A. (1992) Ein Vorkommen foliicoler Flechten in der Steiermark. Herzogia 9: 

239–246. 

Puntillo, D. & Vězda, A. (1994) Some foliicolous lichens new to Calabria. Webbia 49: 125–131. 

Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. W. & Moore, D. M. (1992) The 

Lichen Flora of Great Britain and Ireland. London: Natural History Museum and the British 

Lichen Society. 

Roux, C. & Bricaud, O. (1993) Studo de la genero Strigula (Lichenes, Strigulaceae) en S-Francio. 

Graveco de la makrokonidioj. Bulletin de la Société linnéenne de Provence 44: 117–134. 

Roux, C., Bricaud, O., Sérusiaux, E. & Coste, C. (1994) Wentiomyces lichenicola subsp. nov. 

bouteillei, champignon lichénicole non lichénisé (Dothideales, Dimeriaceae). Mycotaxon 50: 

459–474. 

Santesson, R. (1952) Foliicolous lichens I. A revision of the obligately foliicolous, lichenized 

fungi. Symbolae Botanicae Upsalienses 12(1): 1–590. 

Santesson, R. (1986) Fungi Lichenicoli Exsiccati, fasc. 3–4. Thunbergia 3: 1–18. 

Santesson, R. (1993) The Lichens and Lichenicolous fungi of Sweden and Norway. Lund: SBTförlaget. 

Sérusiaux, E. (1978) Contribution à l’étude des lichens du Kivu (Zaïre), du Rwanda et du 

Burundi. II. Espèces nouvelles de lichens foliicoles. Lejeunia N.S. 90: 1–18. 

Sérusiaux, E. (1985) Goniocysts, goniocystangia and Opegrapha lambinonii and related species. 

Lichenologist 17: 1–25. 

Sérusiaux, E. (1986) The nature and origin of campylidia in lichenized fungi. Lichenologist 18: 

1–35. 

Sérusiaux, E. (1989) Foliicolous lichens: ecological and chorological data. Botanical Journal of the 

Linnean Society 100: 87–96. 

Sérusiaux, E. (1993) New taxa of foliicolous lichens from Western Europe and Macaronesia. 

Nordic Journal of Botany 13: 447–461. 

Sérusiaux, E. (1995) Further new lichen species producing campylidia or complex conidiomata. 

Bibliotheca Lichenologica 58: 411–431. 

Sérusiaux, E. & De Sloover, J. R. (1986) Taxonomical and ecological observations on foliicolous 

lichens in northern Argentina, with notes on the hyphophores of Asterothyriaceae. 

Veröffentlichungen des Geobotanischen Institutes der Eidgenossche Technische Hochschule, Stiftung 

Rübel, Zürich 91: 260–292. 

Tønsberg, T. (1992) The sorediate and isidiate, corticolous crustose lichens in Norway. 

Sommerfeltia 14: 1–331. 

Vězda, A. (1975a) Foliicole Flechten aus der Republik Guinea (W-Afrika). III. Acta Musei 

Silesiae, Opava Ser. A 24: 117–126. 

Vězda, A. (1975b) Foliikole Flechten aus Tanzania (Ost-Afrika). Folia Geobotanica Phytotaxonomica 

(Praha) 10: 383–432. 

Vězda, A. (1980) Foliicole Flechten aus Zaïre. Die Arten der Sammelgattungen Catillaria und 

Bacidia. Folia Geobotanica Phytotaxonomica (Praha) 15: 75–95. 

Vězda, A. (1983) Foliicole Flechten aus der Kolchis (West-Transkaukasien, UdSSR). Folia 

Geobotanica Phytotaxonomica (Praha) 18: 45–70. 

Vězda, A. (1984) Foliikole Flechten der Insel Kuba. Folia Geobotanica Phytotaxonomica (Praha) 

19: 177–210. 

Vězda, A. (1986) Neue Gattungen der Familie Lecideaceae s.lat. (Lichenes). Folia Geobotanica 

Phytotaxonomica (Praha) 21: 199–219. 

Vězda, A. (1987) Foliicole Flechten aus Zaïre (III). Die Gatung Byssoloma Trevisan. Folia 

Geobotanica Phytotaxonomica (Praha) 22: 71–83. 

Vězda, A. (1994) Neue foliicole Flechten II. Nova Hedwigia 58: 123–143. 

Vězda, A. & Poelt, J. (1988) Beiträge zur Kenntnis der Flechtenflora des Himalaya. I. Einige neue 

oder bemerkenswerte gyalectoide und foliicole Flechten. Nova Hedwigia 47: 415–427. 

Accepted for publication 1 October 1995

Foliicolous lichens from Madeira, with the description of a new ...

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?