Joum. Haltori Bot. Lab. No . 65 .' 311-322 (Dec. 1988)

CONTRIBUTIONS TO THE LICHEN FLORA AND LICHEN VEGETATION OF THE CAPE VERDE ISLANDS. VIII. NEW RECORDS OF LICHEN SPECIES ALREADY KNOWN FROM OTHER MACARONESIAN ARCHIPELAGOSI

G ERHARD FOLLMANN AND BRUNO M IES2

INTRODUCTION Until quite recently, only a few notes were available on the composition of the lichen flora of the southernmost Macaronesian archipelago, the Cape Verde Islands (Republica de Cabo Verde), based mostly on more accidental collections and lacking many significant chorological, ecological, and sociological details (Bouly de Lesdain 1935, Cardoso 1915, Crombie 1878, Foll mann 1976, Follmann & KestIer-Merlin 1974, Krog & 0sthagen 1978, Montagne 1860, Stirton 1875, Tavares 1964). Nevertheless, latest systematic studies on the Cape Verdes - including also the smaller, not much visited islands of the group - showed, that the lichen flora of these volcanic and xeric islands in the Atlantic Ocean some 600 km to the west of the African coast (Senegal) is not as poor as could be assumed formerly (Follmann & M ies 1986, 1888a, 1988b, 1988c, Follmann & Sanchez-Pinto 1987). Working-up new collections and older her­ barium specimens, additional lichen taxa have been found to occur on the Cape Verdes and some critical ones could be confirmed, which are of special chorological interest because all but one are already known from other Macaronesian islands. These are briefly documented and discussed here.

MA TERIALS AND METHODS The present results are mainly based on lichen collections made by the authors on the Cape Verde Islands during spring and autumn 1986, preserved now at KOELN (Botanical Institute, University of Cologne, Cologne, Federal Republic of Germany). Supplementary lichen samples have been used from the following herbaria: FR (Botanical Department, Natural History Museum Senckenberg, Francfort, Federal Republic of Germany), LlSC (Botanical Research Centre, Commission fo r Scientific Studies in Oversea Regions, Lisbon, Portugal), LlSU (Botanical Insti tute, University of Lisbon, Lisbon, Portugal), 0 (Botanical Museum, University of Oslo, Oslo, Norway), PC (Cryptogamic Laboratory, National Museum of Natural History, Paris, France), ups (Institute of Systematic Botany, University of Uppsala, Sweden). In case of unnumbered lichen samples, provisional herbarium numbers have been

1 Part 7: G . Follmann & B. Mies. 1988. Contributions to the lichen flora and lichen vegetation of the Cape Yerde Islands. YIf. Some lichen records new to Greater Macaronesia. Cryptogamie, Bryol. Lichenol.: in press. 2 Geobotanical and Phytotaxonomical Study Group, Botanical Institute, University of Cologne, Gyrhofstr. IS, D-5000 Cologne 41, Federal Republic of Germany. 312 Journ. Haltori Bot. Lab. No. 65 198 8 given; these are marked by brackets (e.g., '" [evs-1Il] = Herbarium of the Botanical Museum of the University of Oslo, Cape Verdean lichen collection P. Sunding, revision number 111 noted on the label). The citation of often used synonyms is limited to three. The islands of the Cape Verdes are cited in alphabetical order. The lichen samples have usually been analyzed by light micro­ scopy, spot tests, and thin layer chromatography, exceptionally also by high performance liquid chromatography. In view of the incomplete knowledge of the area limits and ecological requirements of extra-European lichens, a relatively wide chorological and ecological concept is employed. Most pH-values of lichen substrates have been detected on the basis of herbarium material; therefore they may slightly differ from the habitat conditions. Possible sociological affiliations are indicated making use of additional field notes.

COMMENTED INVENTORY OF NEW FINDINGS 1. Acarospora lavicola J. Stein. - Acarosporaceae Syn. Acarospora heu/feriana Koerb. var. massiliensis Harm., Acarospora massiliensis (Harm.) Magn. The crustose, saxicolous, acidophytic to neutrophytic, xerophytic, disjunct Macaronesian-Mediterranean-Sindian species in question has already been collected on the Canary Islands (Champion & Sanchez-Pinto 1978). According to Steiner and Poelt (1984) it is identical with Acarospora massiliensis (Harm.) Magn. cited for the Cape Verde Islands (Fogo, Santo Antiio) by Bou1y de Lesdain (1935). Although never reaching higher degrees of abundance and dominance, on both archipelagos it can be taken most probably for the characteristic species of a pioneer association of open, volcanic rocks in the upper mountain range (Acarosporetum lavicolae Follm. & Mies). Fogo: Exposed, volcanic rocks, northern rim of the main crater upon Monte Vel ha, scattered, 2000 m, NE, pH 7,0; leg. G . Follmann (fII/1 986), KOELN 30 014-30 016.-Exposed, volcanic rocks, south-western rim of the main crater, scattered, 2000 m, NE, pH 6,5 ; leg. G . Follmann (111/1986), KOELN 33024. - Sunny, tufaceous rocks, rim of the main crater south of Chit das Caldeiras, scattered, 2300 m, N, pH 6, 5; leg. B. Mies (X/1986), KOELN CVM-185-e. Santo Antiio: Exposed, volcanic rocks, north-western rim of Cova in the upper part of Ribeira do Paul, scattered, 1300 m, NW, pH 6,0; leg. B. Mies (IX/1986), KOELN CVM-61-f.

2. Acarospora microcarpa (Nyl.) Wedd. - Acarosporaceae Syn. Acarospora schleicheri (Ach.) Mass. var. microcarpa (Nyl.) Oliv., Acarospora tersa (Nyl.) Hue Saxico10us, acidophytic to neutrophytic and xerophytic, this crustose, Medi­ terranean species has also been observed on the Canary Islands (Tavares 1952). On the Cape Verde Islands it can be considered a member of the Macaronesian Acaro­ sporetum lavicolae Follm. & Mies mentioned before. The two samples available for th moment come close to Acarospora microcarpa (Nyl.) Wedd. var. dispersella Magn., <> morphotype without major taxonomic significance. Fogo: Open, volcanic rocks, north-western rim of the main crater upon Monte Velha, rare, 2100 m, NE, pH 6, 5; leg. G . Follmann (111 /1986), KOELN 30021. - Partly wet, tufaceous rocks, inner edge of the main crater west of Chit das Caldeiras, scattered, 1800 m, NE, pH 7,0; leg. B. Mies (X / 1986), KOELN cVM-I 72-k. G . FOLLMANN & B. MIES: Lichen flora and vegetation of the Cape Verde Isls. VIII 313

3. Caloplaca citrina (Hoffm.) T. M. Fries - Teloschistaceae Syn. Blastenia cilrina (Hoffm.) Bouly de Lesd., Pyrenodesmia cilrina (Hoffm.) Trev. This crustose, generally saxicolous, exceptionally corticolous, subneutrophytic to basiphytic, nitrophytic, synanthropic, subcosmopolitan species shows a certain con­ centration in temperate and subtropical regions and has already been noted from the Canary Islands (Tavares 1952). At adequate localities of other areas often appearing in larger populations, here it seems to be a rare inhabitant of eutrophicated rocks in the mountain belt. Fogo: Half-shadowy base of a volcanic boulder in a small vineyard, north-western rim of the main crater upon Monte Vel ha, scattered, 1800 m, SE, pH 7, 0; leg. G. Follmann (1][/1986), KOELN 30027.

4. Cladonia coniocraea (Floerke) Spreng. - Cladoniaceae Syn. Cenomyce coniocraea Floerke, Cladonia jimbriata (L.) E. M. Fries var. coniocraea (Floerke) Vain. The fruticose, humicolous, lignicolous, or terricolous, acidophytic, subcosmo­ politan species cited above has also been observed on the Canary Islands (Champion & Sanchez-Pinto 1978). In Central Europe, for instance, characterizing the widespread humicolous and lignicolous associations of the Cladonion coniocraeae Duvign., on the Canary Islands and still more on the Cape Verdes it seems to play a subordinate role in the ground vegetation of forested parts in the cloud belt. Fogo : Half-shadowy, gravelly loam, together with Cladonia chlorophaea (Floerke) Spreng., aside the defile from Monte Velha to Mosteiros, scattered, 1500 m, NE, pH 7,0; leg. G. Follmann (ITI/1986), KOELN 33027.

5. Endocarpon p:Js/Num Hedw. - Verrucariaceae Syn. Dermatocarpon pusillum (Hedw.) Anzi, Endocarpon glomerulijerum (Mass.) Trev., Endocarpon schaereri (He pp) Fink Normally terricolous, sometimes muscicolous or saxicolous, this squamose, basiphytic, cosmopolitan species has already been cited from Madeira (Tavares 1952). Representing a new genus record for the Cape Verdes, it can be considered a rare rock inhabitant of the up per mountain range for the time being. Fogo: Single, tufaceous boulder, inner slope of the main crater above Chii das Caldeiras, scarce, 1800 m, E, pH7, 0; leg. B. Mies (X/ 1986), KOELN cVM-17 1-h.-Volcanic rocks, ridge of the main crater above Chii das Caldeiras, rare, 2500 m, E, pH 7,0; leg. B. Mies (X/ 1986), KOELN cVM-185-b. Santo Anliio: Terricolous, upper part of Ribeira da Janela, frequent, 1200 m, E, pH 7, 0; leg. B. Mies (IX/ 1986), KOELN cVM-76-i.

6. Fu/gensia subbracteata (Nyl.) Poelt - Teloschistaceae Syn. Caloplaca subbracteata (Nyl.) Lett., Fulgensia sorediosa Klem., Lecanora subbracteata Nyl. This squamose, generally terricolous, exceptionally muscicolous or saxicolous, basiphytic, Mediterranean taxon occurs also on the Canary Islands (Follmann & Poelt 1981) and represents likewise a new genus record for the Cape Verdes. On all three islands cited below only small populations have been observed; therefore it can 314 Journ. Hattori Bot. Lab. No. 65 198 8 pass for a rare ground and rock inhabitant of the mountain range. Fogo: Partly wet soil and tufa, inner slope of the main crater above Cha das Caldeiras, scarce, 1800 m, E, pH 7,0; leg. B. Mies (X/1986), KOELN cVM-l72-c. Santo Antao: Mossy ground, north­ western crater rim of Cova in the upper part of Ribeira do Paul, scarce, 1300 rn, S, pH 6, 5; leg. B. Mies (IX/1986), KOELN cVM-l72-c. Sao Tiago: Volcanic rocks, summit of Pico da Antonia, scarce, 1390 m, W, pH 7, 0 ; leg. B. M ies (X/1986), KOELN cVM-393-g.

7. Lecanora campestris (Schaer.) Hue - Lecanoraceae Syn. Lecanora sub/usca (L.) Ach . var. campestris (Schaer.) Rabenh. Being a variable, crustose, saxicolous, subacidophytic to subbasiphytic, xero­ phytic, synanthropic, Boreomediterranean species, Lecanora campestris (Schaer.) Hue has already been found on the Canary Islands and Madeira (Tavares 1952). On the Cape Verde Islands it can be considered a rare rock inhabitant in the lower mountain range for the moment. Sao Tiago : Volcanic rocks, upper part of Ribeira de Sao Domingos, scarce, 400 m, SE, pH 7, 0 ; leg. L. Sanchez-Pinto (VlII/1978), KOELN 22 138.

8. Lecanora conizaea (Ach.) Nyl. - Lecanoraceae Syn. Lecanora symmicta (Ach.) Ach . var. conizaea (Ach.) Oliv., Lecanora varia (Hoffm.) Ach . var. conizaea (Ach.) Nyl., Rinodina conizaea (Ach.) S. Gray This crustose, corticolous, acidophytic, temperate to subtropical lichen species is also known from the Canaries and Madeira (Tavares 1952). On the Cape Verdes it has been found as a companion species of the local subassociation of the Macaronesian Ramalinetum subgeniculatae Klem. in the lower mountain range. Sao Tiago : On decomposing bark in the dry valley bottom, together with Ramalina subgeniculata Nyl., Ribeira Longueira above Sao Jorge, scattered, 400 m, NE, pH 6, 0 ; leg. C. Brochmann & O. H . Rustan (XIl/1981), 0 (cvB-446-81), KOELN 33020.

9. Lecidea atrobrunnea (Lam. et A. P. de Cand.) Schaer. - Lecideaceae Syn. Lecidea racovitzae Dodge, Psora atrobrunnea (Lam. et A. P. de Cand.) Mass., Rhizocarpon atrobrunneum Lam. et A. P. de Cand. The variable, crustose, saxicolous, acidophytic, subnitrophytic species in question is a subcosmopolitan oreophyte. Cited with some hesitation for the Canary Islands by Tavares (1952), it has recently been confirmed from Tenerife by Crespo and Naranjo (1981). On the Cape Verdes it seems to be one of the rarer pioneer lichens of the upper mountain range, perhaps affiliated with the local subassociation of the Macaronesian Acarosporetum laJlicolae FoIlm. & Mies. Fogo: Volcanic outcrops, together with Acarospora la vicola J. Stein., rim of the main crater upon Monte Vel ha, scattered, 2000 m, NE, pH 6, 5; leg. G . Follmann (IIl/1 986), KOELN 33051 - 33052.

10. Ochrolechia pallescens (L.) Mass. - Lecanoraceae Syn. Lecanora pal/escens (L.) Ach., Ochrolechia parella (L.) Mass. var. pal/escens (L.) Rabenh., Pertusaria pallescens (L.) M uell.-Arg. Strictly corticolous, subacidophytic to subneutrophytic, this subcosmopolitan, G. FOLLMANN & B. MIEs: Lichen flora and vegetation of the Cape Verde Isls. VIII 315 crustose lichen shows suboceanic distribution tendencies. Already known from the Canaries and Madeira (Tavares 1952), together with the following it represents a new genus record for the Cape Verdes. Actually it can be taken for a rare companion species of the local subassociation of the Macaronesian Ramalinetum subgeniculatae Klem. Sao Tiago : Branches of Ceratonia siliqua L., together with Ramalina subgeniculata Nyl., Ribeira de Sao Jorge, scarce, 750 m, SE, pH 6,0; leg. B. Mies (X/1986), KOELN cVM-236-e.

11. Ochrolechia parella (L.) Mass. - Lecanoraceae Syn. Lecanora parella (L.) Ach., Ochrolechia pallescens (L.) Mass. var. parella (L.) Koerb., Per­ tusaria incarnafa Leight. This crustose, acidophytic to subneutrophytic, Subatlantic species, the saxicolous counterpart of the preceding, occupies a Central European to Mediterranean area and has so far been reported from the Azores, Canaries, and Madeira (Tavares 1952). Although well fruiting and rich in variolaric acid, both samples cited below exhibit comparatively small and thin thalli coming close to Ochrolechia parella (L.) Mass. f. tenuis Vers., a morphotype of subordinate taxonomic significance. For the moment it can pass for one of the rarer pioneer lichens on hard volcanic rocks in the mountain range of the Cape Verdes. Boa Vista: Smooth basaltic rocks, ascent to Monte Estancia, scarce, 300 m, SE, pH 6, 5; leg. W. Lobin (IX/1979), FR CVL-380, KOELN 33021. Sao Vicente: Smooth basaltic rocks, summit of Monte Verde, scarce, 700 m, NW, pH 6, 0; leg. L. Sanchez-Pinto (VIII/l978), KOELN 33022.

12. Phaeophyscia hispidula (Ach.) Mob. - Physciaceae Syn. Parmelia hispidula Ach., Physcia hispidula (Ach.) Frey, Physcia setosa (Ach.) Frey Being a variable, normally corticolous, rarely saxicolous, subacidophytic to subneutrophytic, oceanic, temperate to tropical phyllolichen, Phaeophyscia hispidula (Ach.) Mob. has hitherto been recorded for the Canary Islands and Madeira (Tavares 1952). On the Cape Verdes it can actually be considered to be a rare epiphyte and lithophyte of the mountain range. Santo Antao: Loamy rock face, north-west of Cova in the upper part of Ribeira do Paul, scattered, 1130 m, NW, pH 7, 0; leg. P. Sunding (X/ 1972), 0 (cvs-414). - Branches of Euphorbia tuckeyana Steud., together with Heterodermia leucomela (L.) Poelt and Parmelia fiaventior Stirt., near pass be­ tween Porto Novo and Ribeira Grande, scarce, about 1000 m, NW, pH 6, 5; leg. K . Lewejohann (XH/l978), KOELN 22 121.

13 . Physcia dimidiata (Am.) Nyl. - Physciaceae Syn. Parmelia dimidiata (Am.) Am., Physcia aipolia (Humb.) Fuern. var. dimidiata (Am.) Nyl., Physcia albinea (Ach.) Malbr. var. dimidiata (Am.) Nyl. This foliose, corticolous or saxicolous, subneutrophytic to subacidophytic, Central European to Mediterranean taxon related to Physcia dubia (Hoffm.) Lett. has already been cited for the Canary Islands by Tavares (1952). In view of the single sample avail­ able for the moment it seems to be a rare rock inhabitant of the cloud belt of the Cape Verdes. It should be noted that our specimen clearly does not belong to the morpholo­ gically deviating Physcia dimidiata (Am.) Nyl. var. ornata (Nadv.) Mob. reported from 316 Journ. Hattori Bot. Lab. No. 65 198 8

Africa by Moberg (1986). Sal: Volcanic rocks, summit of Monte Grande, scarce, 400 m, NE, pH 7, 0; leg. P. Sunding (XI 1972),0 (cvs-424).

14. Pyxine sorediata (Ach.) Mont. - Physciaceae Syn. Lecidea sorediata Ach ., Pyxine cocoes (Swartz) Nyl. var. sorediata (Ach.) Tuck., Physcia endochrysea Nyl. Sometimes corticolous, sometimes saxicolous, the foliose, acidophytic species in question is widespread in temperate and tropical regions. It has been noted for the Azores by Degelius (1941), for the Canaries by Klement (1965), and for Madeira by Arvidsson and Wall (1985). Like Pyxine cocoes (Swartz) Nyl. (Follmann & Sanchez­ Pinto 1987) Pyxine sorediata (Ach.) Mont. represents a relatively common epiphyte and lithophyte in the mountain range of the Cape Verdes. Brava : Exposed, volcanic boulder, south-west of Fundo Grande, scarce, 800 m, N , pH 6, 5; leg. A. Kalnins (XI/ 1986), KOELN cVM-4 16-a. Santa Antao: Volcanic rocks, carriage road from Paul to Cova, abundant, 1200 m, NE, pH 6,0; leg. K . Bys trom (XI/1958), L1SU cVT-8 1. Sao Nicolau : Stem of Dracaena draco (L.) L., Cruzetinha, scattered, 850 m, NW, pH 6, 0; leg. K. Bystrom (V/1959), L1SU cVT-970. Sao Tiago : Branches of Euphorbia tuckeyana Steud., south-western slope of Pico da Antonia, scarce, 1250m, SW, pH 7, 0 ; leg. B. Mies (X/ 1986), KOELN cVM-388 -m. - Tufaceous rocks, steep slope in the Serra da Antonia, abundant, 1100 m, NE, pH 6,5 ; leg. B. Mies (X/1986), KOELN cVM-223-a. - Volcanic rocks, near summit of Pico da Antonia, scarce, .1350 m, E, pH 6, 5; leg. B. Mies (X/1986), KOELN cVM-390-c.

15. Ramalina canariensis J. Stein. - Ramalinaceae This foliose, corticolous, acidophytic, Macaronesian-Mediterranean species has originally been described from the Canary Islands, is registered by Tavares (1952) fo r the Azores, and could recently also be discovered on Madeira (Follmann unpubl.). On the Canaries being a characteristic species of the local subassociation of the Macaronesian Ramalinetum subgeniculatae Klem., on the Cape Verdes it seems to be an extremely rare epiphyte in the lower mountain range. Santa Antao: Rotten wood, central basin of Ribeira Grande, scarce, 150 m, W, pH 6,0; leg. G. Follmann (IVj I974), KOELN 28 795.

16. Ramalinafarinacea (L.) Ach. - Ramalinaceae Syn. Ramalina /allax Mot., Ramalina motykana Bystr., Ramalina reagens (Bouly de Lesd.) W. L. Culb. Recorded for all Macaronesian archipelagos (Tavares 1952), this polymorphic, fruticose, generally corticolous, exceptionally saxicolous, subacidophytic to subneutro­ phytic, North Hemispheric entity has supposedly a Macaronesian-Mediterranean distribution centre (Krog & 0sthagen 1980). The first report from the Ca pe Verde Islands is Stirton's (1875), except the hint "supra terram" unfortunately lacking all other collection data. In his revision of the lichen collections of the Challenger Expedi­ tion, Crombie (1878) pointed out that the sample seen by Stirton belongs to Ramalina intermedia (Del.) Nyl. Also Cardoso's (1915) collections from Santo Antao and Sao G . FOLLMANN & B. MIES : Lichen flora and vegetation of the Cape Verde Isls. VIII 317

Nicolau are said to correspond to this chorologically and ecologically deviating taxon often confused with Ramalina subfarinacea (Nyl.) Nyl., an opinion shared by Bouly de Lesdain (I 935). Ramalinafarinacea (L.) Ach. is confirmed here for the Cape Verde Islands, where it occurs as a locally abundant companion species of various epiphytic lichen associa­ tions at mountain sites with a humid microclimate. The statement "The Canary Islands may constitute its southern most limit in the Atlantic region" (Krog & 0sthagen 1980) should therefore be rectified. All intermediate stages between fine-lobed and broad­ lobed morphotypes (var. gracilenta Ach. - var. pendulina Ach.) and also some fertile thalli have been found among the samples cited below. To judge from the spot tests, these represent various uncorrelated chemotypes, the acid-deficient one being the most frequent. According to our recent observations, the mainly Nearctic Ramalina intermedia (Del.) Nyl. is absent from the Cape Verdes, and its existence on the other Macaronesian islands seems to be highly questionable. Brava: Bark of cultivated trees, together with Ramalina peruviana Ach., Nova Sintra, frequent, 500 rn , NE, pH 6, 0 ; leg. A. Kalnins (X/ 1986), KOELN cVM-408-a. - Epiphytic, Cova Ribeira da Santa Cruz south-west of Nova Sintra, abundant, 500 rn , NE, pH 6, 0; leg. A. Kalnins (X/1986), KOELN cVM-407-a. - Dry twigs of deciduous trees, Monte Mato south of Nova Sintra, abundant, 600 rn , NE, pH 6, 5; leg. A. Kalnins (XI/ 1986), KOELN cVM-41O-b. Fogo: Epiphytic, Matinho near Mosteiros, abundant, 1000 rn , NE, pH 6,0; leg. A. Chevalier (lX/1934), PC (cvc-2682). Samo Antao : Epiphytic, north-east of Cova in the upper part of Ribeira do Pa(tl , scarce, 1150 rn , NE, pH 6, 0; leg. L. Borgen (1 / 1980),0 (c vB-1 13). Sao Nicolau : Dry shrubs, north-eastern slope of Monte Agua de Jose, scattered, 650m, NE, pH6, 5; leg. G. Follrnann (Ill/ I 986), KOELN 33133. Sao Tiago: Bark of Faidherbia albida (Del.) Chev., upper part of Val de Sao lorge dos Orgaos, abundant, 800 rn, NE, pH 6, 0; leg. G . Follrnann (IV/1974), KOE LN 33 134. - Dry twigs of Ceratonia siliqua L., upper part of Val de Sao Jorge dos Orgaos, abundant, 700 rn , SE, pH 6, 0; leg. B. Mies (X/1986), KOELN cVM-277-b. - Bark of Ziziphus jujuba Mil l. , upper part of Val de Sao Jorge dos Orgaos, abundant, 880 rn, E, pH 6, 0 ; leg. B. Mies (X/1986), KOELN cVM-229-g.

17 . Ramalina huei Harm. - Ramalinaceae Syn . Ramalina dracaena Magn., Ramalina superfraxinea Follrn. & Sanch.-Pinto Till now this fruticose, corticolous, acidophytic species has been taken for a local Canarian endemic (Krog & 0sthagen 1980). Contrary to most Canarian collections, our single Cape Verdean sample is comparatively small, but corresponds well in secondary chemistry (divaricatic and usnic acids, triterpenoids) and anatomical struc­ ture. Like on the Canaries it seems to be sociologically related to the local subassocia­ tion of the Macaronesian Ramalinetum subgeniculatae Klem. Sao Tiago : Bark of Eucalyptus globulus Lab., upper part of Ribeira da Janela between Rui Vaz and Sao Jorge, scarce, 680 m, NE, pH 6, 5; leg. L. Borgen (1 /1 980), 0 (cvM-114).

18. Ramalina subfarinacea (Nyl.) Nyl. - Ramalinaceae Syn. Ramalina angustissima (Anzi) Vain., Ramalina farinacea (L.) Ach. var. angustissima Anzi, Ramalina scopu/orum (Retz.) Ach. var. subfarinacea Nyl. Ramalina subfarinacea (Nyl.) Nyl. externally resembles Ramalina farinacea (L.) Ach., differing nevertheless in morphological (spiky appearance, spreading hold fast, 318 Journ. Hattori Bot. Lab. No. 65 198 8 excavate soralia), ecological (coastal rocks), and chorological characteristics (American­ European distribution area). Although registered from all other Macaronesian archi­ pelagos (Tavares 1952), it could not be refound on the Canary Islands by Krog & 0sthagen (\980). Our single Cape Verdean specimen, associated with Teloschistes flavicans (Swartz) Norm., belongs to the predominant chemotype with norstictic and usnic acids (Krog & lames 1977). Santo Antao: Volcanic rocks, together with Teloschistes flavicans (Swartz) Norm., upper part of Ribeira Cha da Pedra, scattered, 820-900 m, NE, pH 6, 0 ; leg. P. Sunding (X[/ 1976), 0 (evs-112).

19. Roccella linearis (Ach.) Vain. - Roccellaceae Syn. Roccella /uci/ormis (L.) A. P. de Cand. var. linearis Ach., Nemaria /uci/ormis (L.) Nav. var. linearis (Ach.) Nav. Sometimes corticolous, sometimes saxicolous, but always acidophytic and halo­ phytic, this fruticose species related to Roccella montagnei Bel. inhabits a large, dis­ junct Macaronesian-Paleotropical area and is already known from the Azores and Madeira (Tavares 1952). Due to the thallus reactions (C+ red, K + pink) our samples belong formally to Roccella linearis var. guineensis Vain., but it is not yet clear whether the different varieties described by Vainio (1901) are mere chemotypes or not. Un­ doubtedly having been a companion species of the Cape Verdean subassociation of the Macaronesian Roccelletum canariensis Follm. of the coastal belt, it could not be refound in recent times and might actually be extremely rare or extinct due to over­ collecting for commercial purposes. Ilhas de Cabo Verde: Probably on volcanic rocks, together with Roccella belangeriana Awas. , without further collection data ; certainly before 1896, Herb. GiUet s. n., ups s. n. Sao Nicolau: Volcanic rocks, together with Roccella montagnei Bel. and Roccella vicentina (Vain.) Follm., Punta do Sal, probably near Cha da Boca da Ribeira, without further collection data; LIse 30-A-1.

20. Thermutis velutina (Ach.) Flot. - Lichinaceae Syn. Gonionema velutinum (Ach.) Nyl., Polychidium vetutinum (Ach.) S. Gray, Scytonema velutillum (Ach.) Wallr. This primitive, filiform, saxicolous, relatively euryecious species covers a wide, but discontinuous Boreal area. Apparently absent from the other Macaronesian archi­ pelagos, until now it has only been cited once from a small rocky islet off the north­ western coast of Sao Vicente (Ilheu dos Passaros: Stirton 1875), an assertion ques­ tioned by Crombie (1878) and Tavares (1952). Its existence on the Cape Verde Islands is confirmed here, where it occurs sporadically - mostly together with cyanobaeteria and bryophytes - on temporary wet, rapidly drying-up, half-shadowy, volcanic rocks in the mountain range. Santo Antao: Between mosses on wet, volcanic rocks, south-eastern part of the inner crater rim of Cova in the upper part of Ribeira do Paul, scattered, 1150 m, NW, pH 6, 0; leg. B. Mies (IX/ I 986), KOELN eVM-63-v. - Between mosses on wet, volcanic rocks, north-western crater rim of Cova in the upper part of Ribeira do Paul, scattered, 1300-1499 m, NE, pH 6,0; leg. B. Mies (IX/1986), KOELN eVM-61 -k. Sao Tiago: Wet, tufaceous rocks, steep slope in the Serra da Antonia, scarce, 1l00m, NE, pH 6,0; leg. B. Mies (X/ l 986), KOELN eVM-223-b. - Wet, tufaceous rocks, Val de Picos at the G . FOLLMANN & B. MIES : Lichen flora and vegetation of the Cape Verde Isls. VIn 319

northern slope of Serra da Antonia, scarce, 990 m, NW, pH 6, 0; leg. B. Mies (X/1986), KOELN CVM- 220-e.

21. Trape/ia eoaretata (l. E. Smith & Sower.) Choisy - Trapeliaceae Syn. Biatora coarctata (J. E. Smith & Sower.) T. M. Fries, Lecanora coarctata (J. E. Smith & Sower.) Ach., Lecidea coarctata (J. E. Smith & Sower.) Nyl. The crustose, saxicolous or terricolous, acidophytic, subcosmopolitan species in question shows a certain preference for Mediterranean regions. Already reported from the Canaries (Champion & Sanchez-Pinto 1978) and Madeira (Tavares 1952), it represents a new genus record for the Cape Verdes, where it can be considered a rare lithophyte of the mountain range for the moment. Fogo: Fissures of tufaceous rocks, Ribeira Tlheu, scarce, 700-900 m, N, pH 6, 0; leg. B. Mies (X/1986), KOELN CVM-149-h. Santo Antao : Volcanic rocks, crater of Cova in the upper part of Ribeira do Paul, scarce, 1150 m, NE, pH 6, 0 ; leg. K . Bystrom (X/ 1958), LISU CVT-147.

22. Usneafragileseens Hav. - Usneaceae Usnea fragislescens Hav., a preferably saxicolous, acidophytic, Atlantic-European species, is known as yet from the Canary Islands (Champion & Sanchez-Pinto 1978). Contrary to the related Usnea mollis Stirt., reported recently from forest trees on the Cape Verdes (Follmann & Mies 1988a), it seems to be affiliated with the epilithic variant of the pantropical Telosehistetum flavieantis Follm. in the mountain range. Brava : Volcanic cliffs, east-northeastern slope of Fetos above Ribeira da Gan;a, scattered, 760 m, NE, pH 6, 0; leg. O. H. Rustan & c. Brochmann (1II/1982), 0 (cvR-258). Santo Antao: Volcanic rocks, together with Heterodermia boryi (Fee) Hale, north-west of Cova in the upper part of Ribeira do Paul, scattered, 1240 m, NE, pH 6, 0 ; leg. P. Sunding (X/ 1972), 0 (cvs-260). - Volcanic rocks, together with Teloschistes flavicans (Swartz) Norm., northeast of Cova in the upper part of Ribeira do Paul, scattered, 1080 m, NW, pH 6, 0; leg. L. Borgen (1 /1980), 0 (cvB-257, cVB-259, CVB-261) . Siio Tiago: Stalks of Euphorbia luckeyana Steud., western slope near summit of Pico da Antonia, scarce, 1300 m, S. pH 6, 0; leg. B. Mies (X/1986), KOELN cVM-388 -b.

23. Xanthoparmelia subramigera (Gyeln.) Hale - Parmeliaceae Syn. Parmelia canariensis Gyeln., Parmelia lavicola Gyeln., Parmelia subramigera Gyeln. Originally saxicolous, sometimes "subterricolous", acidophytic to subneutrophytic, this variable, foliose species represents one of the commonest members of the genus in tropical and subtropical regions, extending here and there into temperate zones, and already known to occur on the Canary Islands (Champion & Sanchez-Pinto 1978). The samples cited below consist of broad- and fine-lobed, even and curled, moderately and intensely isidiose morphotypes, some of which could have been taken by Bouly de Lesdain (1935) for Parmelia verrucigera Nyl. (Parmelia eonspersa [Ehrh.] Ach. var. verrucigera [Nyl.] Boist.) mentioned from Fogo. On Santo Antiio Xanthoparmelia subramigera (Gyeln.) Hale seems to be a relatively frequent companion species of the Cape Verdean subassociation of the pantropicaJ Parmotremetum tinetorum Follm. in the mountain range. Santo Antao: Soil-incrusted, volcanic rocks, Lombo Cebide Vila south of Ribeira Grande, scat- 320 Journ. Hattori Bot. Lab. No. 65 198 8

tered, 1100 rn, SE, pH 6, 5; leg. P. Sunding (X/1972), 0 (cvs-34 I , cvs-343). - Open, volcanic rocks, north-west of Cova in the upper part of R ibeira do Paul, scattered, 1130 rn, NE, pH 6, 0; leg. P. Sunding (X/ 1972), 0 (cvs-345, cvs-346). - Open, volcanic rocks, upper part of Ribeira Chit da Pedra, scattered, NE, pH 6, 0; leg. P. Sunding (XI/1976), 0 (cvs-342).

24. Xanthoria resendei Poelt & Tav. - Teloschistaceae Syn. Xanthoria subelegans Tav. & Poelt This subfoliose, saxicolous, subacidophytic to subneutrophytic, Atlantic-Medi­ terranean taxon has been collected so far on the Azores, Canaries, and Madeira (Poelt & Tavares 1968). Contrary to the central and northern Macaronesian islands and southwestern Europe, where it constitutes a locally abundant, maritime li thophyte, on the Cape Verdes it seems to be a rare montane one. Most probably it occurs not only on Sao Tiago, but also on Santo Antao, where its growing site proved to be com­ pletely inaccessible. saG Tiago : Exposed, volcanic boulder, south-western slope of Pico da Antonia, scarce, 1220 rn, N, pH 6, 5; leg. B. Mies (X/ 1986), KOELN cVM-386-a.

DISCUSSION The biogeographical concept of Macaronesia, established by Engler (1879) for "Lauromacaronesia" (Azores, Canaries, Madeira) and used by Bramwell (1972), Knapp (1973), Sunding (1979), and many others for the Cape Verde Islands as well ("Greater Macaronesia"), has recently been criticized from tloristical and faunistical points of view (Claussen & Barkemeyer 1987, Geisthardt 1982, Lobin 1982, etc.). In this context it should be underlined that all but one (Thermutis velutina [Ach.] Flot.) lichen species enumerated before appear also on other - often all - Macaronesian islands. This corresponds we ll to previous observations and points again to close fioristical relations between the four Midatlantic archipelagos, which passed a similar Tertiary history of origin and rise. According to a new check-list of Cape Verdean lichens actually in preparation, roughly 70 % show Cape Verdean-Central Macaronesian occurrences. These tloristical affinities are retlected sociologically by the existence of ma ny Macaronesian lichen-dominated plant associations forming characteristic local sub­ associations on the various archipelagos including the Cape Verdes. Typical examples for those joint lichen communities mentioned in the present inventory are the epilithic, xerophytic Acarosporetum lavicolae Follm. & Mies of the higher mountain range, the epiphytic, aerohygrophytic Ramalinetum subgeniculatae Klem. of the lower mountain range, or the epilithic, halohygrophytic Roccelletum canariensis Follm. of the coastal range. On the other side it appears to be noticeable that only one of the lichen taxa treated can pass for a true Macaronesian endemic (Ramalina huei Harm.), whereas the others show Atlantic-Mediterranean (including Macaronesian-Mediterranean, 8= 33 %; e.g., Xanthoria resendei Poelt & Tav.), Cosmopolitan (including Subcosmo­ politan, 6= 25 %; e.g., Cladonia coniocraea [Floerke] Spreng.), Tropical (including Paleotropical and Subtropical, 5= 21 %; e.g., Xanthoparmelia subramigera [Gyeln.] G. FOLLMANN & B. MI ES: Lichen flora and vegetation of the Cape Verde Is1s . VIII 321

Hale), and Boreal (including Boreomediterranean, 4= 17 %; e.g., Lecanora campestris [Schaer.] Hue) distribution tendencies.

ACKNOWLEDGMENTS The authors are much indepted to the directors and curators of the herbaria at Francfort (FR), Lisbon (usc, usu), Oslo (0), Paris (pc), and Uppsala (ups) for the loan of Cape Verdean lichen samples in their care and the patience they had regarding the return of these. Moreover, they gratefully acknowledge the assignment of lichen collections from the Island of Brava by A. Kalnins (Warendorf) and the confirmations, determinations, and suggestions by D r. K. Kalb (Neumarkt), Dr. S. Kurokawa (Ibaraki), Dr. R. Moberg (Uppsala), and Prof. Dr. J. Poelt (Graz).

SUMMARY Twenty-four lichen species already known from different Macaronesian archipelagos are reported from the Cape Verde Islands and circumscribed chorologically, ecologically, and - if possible- sociolo­ gically. Endoearpon pusillum Hedw., Fulgensia subbraeleala (Nyl.) Poelt, Ochrolechia pallescens (L.) Mass., Ochrolechia parella (L.) Mass., and Trapelia eoare/ala (J.E. Smith & Sower.) Choisy represent new genus records for the Cape Verdes. The existence of Ramalina farinacea (L.) Ach. , Ramalina subfarinacea (Nyl.) Nyl., and ThermUlis velu/ina (Ach.) Flot. on the Cape Verdes is confirmed. Roccella linearis (Ach.) Vain. seems to be extinct on the archipelago due to overexploitation nowadays. Rama­ lina inlermedia (Del.) Nyl. has to be excluded from the catalogue of Cape Verdean lichens. Without detriment to their Macaronesian occurrence, the species treated show the following general distribution tendencies: Atlantic-Mediterranean (including Macaronesian-Mediterranean, 33 %), Cosmopolitan (including Subcosmopolitan, 25 %), Tropical (including Paleotropical and Subtropical, 21 %), Boreal (including Boreomediterranean, 17 %), Macaronesian (4 %).

REFERENCES Arvidsson, L. & S. Wall. 1985. Contribution to the lichen flora of Madeira. Lichenologist 17: 19-49. Bouly de Lesdain, M. 1935. Lichens des lIes du Cap Vert recueillis par A. Chevalier. Rev. bryol. lichenol., N. S. 8 : 14--17. Bramwell, D. 1972. Endemism in the flora of the Canary Islands. In D. H . Valentine (ed.), Taxonomy, phytogeography, and evolution: 141 -159. London. Cardoso, J. 1915. Cryptogamicas das I1has de Cabo Verde. V. Lichenes. Broteria, ser. Bot. 13(2) : 116-123. Champion, C. L. & L. Sanchez-Pinto. 1978. Catalogo preliminar de los liquenes de las Islas Canarias. Santa Cruz de Tenerife. Claussen, C. & W. Barkemeyer. 1987. Zur Syrphidenfauna der Kapverden (Insecta: Diptera : Syrphidae). Cour. Forschungsinst. Senckenberg 95 : 71-86. Crespo, A. & J. Naranjo. 1981. Sobre la flora Iiquenica del Parque Nacional de las Canadas del Teide (Tenerife, Canarias). Lazaroa 3 : 375- 377. Crombie, J. M. 1878. The lichens of the "Challenger Expedition". J. Linn. Soc. London, Bot. 16 : 210-231. Degelius, G. 1941. Lichens from the Azores, mainly collected by H . Persson. Goteborgs kongl. Vetensk. Vitterh. Samh. Handl., ser. B, VU, 1(7) : 3-46. Engler, A. 1879. Versuch einer Entwicklungsgeschichte der extratropischen Florengebiete der nord­ lichen Hemisphiire. Leipzig. 322 Journ. Hattori Bot. Lab. No. 65 198 8

Follmann, G. 1976. Observaciones acerca de la flora y vegetaci6n de liquenes de las Islas de Cabo Verde. 11. Lecanora sulphurella Hepp (Lecanoraceae). Cuad. bot. Canar. 26-27: 1- 7. --- & J.-R. Kestler-Merlin. 1974. Observaciones acerca de la flora y vegetaci6n de liquenes de las Islas de Cabo Verde. I. Pseudocyphellaria aurata (Ach.) Vain. (Stictaceae). Cuad. bot. Canar. 22: 19-21. -- & B. Mies. 1986. Contributions to the lichen flora and vegetation of the Cape Verde Islands. IV. New lichen records and their chorological significance. J. Hattori bot. Lab. 61: 499-523. - - - & - --. 1988a. Zur Kenntnis der Flechtenflora und Flechtenvegetation der Kapverdischen Inseln. V. Neue Hygroskiophyten des Nebelglirtels und ihre Vergesellschaftung. Cour. For­ schungsinst. Senckenberg (in press). --- & --. 1988b. Zur Kenntnis der Flechtenflora und Flechtenvegetation der Kapverdischen Inseln. VI. Zusammenstellung von den Kapverdischen Inseln erstmalig beschriebener Flech­ tensippen (Lichenophyta). Cour. Forschungsinst. Senckenberg (in press). --- & --. 1988c. Contributions to the lichen flora and lichen vegetation of the Cape Verde Islands. VH. Some lichen records new to Greater Macaronesia. Cryptogamie, Bryol. Lichenol. (in press). --- & J. Poelt. 1981. Zur Kenntnis der Flechtenflora und Flechtenvegetation der Kanarischen Inseln. V. Eine neue Bodenflechte aus dem Verwandtschaftskreis von Fulgensia desertorum (Tom.) Poelt und das Diploschistetum albescentis Klem. Philippia 4: 370-378. -- & L. Sanchez-Pinto. 1987. Zur Kenntnis der Flechtenflora und Flechtenvegetation der Kap­ verdischen Inseln. HI. Neue Flechtenfunde und ihre Bedeutung flir die Floren- und Vegetations­ geschichte. Cour. Forschungsinst. Senckenberg 95: 175-188. Geisthardt, M. 1982. Beitrag zur Kenntnis der Kiiferfauna der Kapverden mit zoogeographischen Anmerkungen. Cour. Forschungsinst. Senckenberg 52: 167-171. Klement, O. 1965. Zur Kenntnis der Flechtenvegetation der Kanarischen Inseln. Nova Hedwigia 9: 503-582. Knapp, R. 1973. Vegetationsmonographien der einzelnen Gro13riiume. HI. Die Vegetation von Afrika. Stuttgart. Krog, H. & P. W. James. 1977. The genus Ramalina in Fennoscandia and the British Isles. Norw. J. Bot. 24 : 15- 43. --- & H. 0sthagen. 1978. Three new Ramalina species from Macaronesia. Norw. J. Bot. 25: 55-59. -- & --. 1980. The genus Ramalina in the Canary Islands. Norw. J. Bot. 27: 255- 296. Lobin, W. 1982. Untersuchungen liber Flora, Vegetation und biogeographische Beziehungen der Kapverdischen Inseln. Cour. Forschungsinst. Senckenberg 53: 1-112. Moberg, R . 1986. The genus Physcia in East Africa. Nord. J. Bot. 6: 843-864. Montagne, C. 1860. Florula gorgonea seu enumeratio plantarum cellularium, quas in Promontorio Viridi (Cap Vert) insuliisque adjacentibus a diversis botanicis et imprimis C. Bolle, berolinensi, hucusque collectas, recognovit descripsitque. Ann. Sci. nat. (Paris), ser. IV, 14: 210-225. Poelt, J. & c. N. Tavares. 1968. Xanthoria resendei, eine neue Laubflechte der siidwesteuropiiischen Kiisten. Port. Acta bioI., ser. B, 9: 300-307. Steiner, M. & J. Poelt. 1984. Flechten aus Afghanistan. 1. Acarospora subgen. Acarospora. Nova Hedwigia 39: 559-567. Stirton, J. 1875. Enumeration of the lichens collected by H . N . Moseley, M. A., naturalist to H . M. S. "Challenger", in the islands of the Atlantic Ocean. J. Linn. Soc. London, Bot. 14: 366-375. Sunding, P. 1979. Origins of the Macaronesian flora. In D . Bramwell (ed.), Plants and islands: 13-40. London. Tavares, C. N. 1952. Contributions to the lichen flora of Macaronesia. I. Lichens from Madeira. Port. Acta bioI., ser. B, 3: 308-391. - --. 1964. Contributions to the lichen flora of Macaronesia. Ill. New or interesting taxa. Rev BioI. (Lisboa) 4: 131-144.