CONTRIBUTIONS TO THE LICHEN FLORA AND ... - J-Stage

Joum. Hattori Bot. Lab. No. 61 : 499-523 (Dec. 1986)

CONTRIBUTIONS TO THE LICHEN FLORA AND LICHEN VEGETATION OF THE CAPE VERDE ISLANDS. IV.

NEW LICHEN RECORDS AND THEIR CHOROLOGICAL SIGNIFICANCP

GERHARD FOLLMANN AND BRUNO MIES2

INTRODUCTION

Although pioneer lichens play an important role in the colonization of younger lava flows and characterize large parts of the landscape of the Cape Verde Islands, little is known about their taxonomy, chorology, ecology, and sociology compared with the other Macaronesian archipelagos (Azores, Canaries, Madeira). Based on older and sometimes dubious records, Tavares (1952) listed about 90 lichen taxa, being less than 10% of the lichen flora of the entire Macaronesian region known at that time. Including the observations of Follmann and Sanchez-Pinto (1986), the lichen inventory of the Cape Verdes actually comprises 177 items. Nevertheless, deduced from larger collections made during the last decade, especially of hitherto neglected microlichens, the total number oscillates probably between 300 and 400. A series of further new records for Macaronesia, the Cape Verdes, or single islands of the latter is briefly documented and discussed here.

MATERIALS AND METHODS

The present results are mainly based on lichen collections made on the Cape Verde Islands by Sunding (1972, 1976), Follmann (1974, 1986), Lobin (1978, 1979), Sanchez-Pinto (1978), Borgen (1980), and Rustan and Brochmann (1981, 1982). Voucher specimens are preserved at FR (Natural History Museum "Senckenberg", Francfort, Federal Republic of Germany: Lobin), KASSEL (Natural History Museum "Ottoneum", Kassel, Federal Republic of Germany: Follmann), KOELN (Botanical Institute, University of Cologne, Cologne, Federal Republic of Germany: Follmann, Lobin, Sanchez-Pinto), 0 (Botanical Museum, University of Oslo, Oslo, Norway : Borgen, Rustan and Brochmann, Sunding), and TFMC

(Natural History Museum, Santa Cruz, Tenerife, Canary Islands: Sanchez-Pinto). Since the lichen samples from 0 and TFMC remained unnumbered, provisional herbarium numbers are given in brackets. The citation of often used synonyms is limited to three. Some of the more abundant species have been (or will be) distributed with the Lichenes Exsiccati Selecti a Museo Historiae Naturalis Casse1ensi Editi (LESK) and the Lichenes Exsiccati Selecti ab Instituto Botanico Universitatis Coloniensis Editi (LESC).

1 Part 3: Follmann, G. and Sanchez-Pinto, L., 1986. Zur Kenntnis der Flechtenflora und Flechtenvegetation der Kapverdischen Inseln. Ill. Neue Flechtenfunde und ihre Bedeutung fUr die Florenund Vegetationsgeschichte. Cour. Forschungsinst. Senckenberg 86: in press.

2 Botanical Institute, University of Cologne, Gyrhofstr. IS, D-5000 Cologne 41 , Federal Republic of Germany.

500

17°

Journ. Hattori Bot. Lab. No. 61

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<6Sanla Luzia

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ILHAS DE CABO VERDE

1 9 8 6

16°--------+----------------------r--------------------~~~~----~

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FIG. I. General situation and highest elevations of the Cape Verde Islands.

Most pH-values of lichen substrata have been detected on the basis of herbarium material. If at all possible, specific secondary lichen products have been determined by thin layer chromatography (TLC) and high performance liquid chromatography (HPLC). In view of the incomplete knowledge of the area limits of extra-European lichens, a relatively wide chorological concept is employed for the phytogeographical arrangemen.t of the new records: Cosmopolitan geoelements (including Subcosmopolitan and disjunct ones), Tropical geoelements (including Neotropical, Paleotropical, and Subtropical ones), Mediterranean geoelements (including Atlantic-Mediterranean and Macaronesian-Mediterranean ones), and Macaronesian geoelements (including local endemics). The principal ecotypes are treated in a similar manner: epiphytes, geophytes, and lithophytes; acidophytes, basiphytes, and neutrophytes; xerophytes, mesophytes, and hygrophytes. General informations on origin, terrain, climate, flora, and vegetation of the Cape Verde Islands are given by Chevalier (1935: botany), FriedHinder (1913 : geology), Hansen and Sunding (1985: floristics), Klug (1973: geography, 1980: climatology), Lobin (1982: chorology), Machado (1965 : vulcanism), Rothe (1982: geology), and Sunding (1973: endemism, 1977 : bibliography).

G. FOLLMANN & B. MIES: Lichen flora and vegetation of Cape Verde Islands. IV 501

RESULTS

COSMOPOLITAN GEOELEMENTS

I. Catapyrenium lachneum (Ach.) Sant. - Verrucariaceae Syn. Dermatocarpon hepaticum (Ach.) T. M. Fries, Dermatocarpon lachneum (Ach.) A. L. Smith, Dermatocarpon rufescens (Ach.) T. M. Fries. This placodioid, generally terricolous, occasionally saxicolous, basiphytic to

neutrophytic, Subcosmopolitan pyrenolichen has hitherto been reported from the Canary Islands (Topham & Swinscow 1982) and Madeira (Tavares 1952). It is new to the Cape Verde Islands. Although the only specimen seen is well developed, it must be considered a rare soil inhabitant in the mountain belt at present.

Sao Nicolau: Sandy loam, summit of Monte Gordo, scarce, 1200 m, NE, pH 7,0; leg. W. Lobin (XII/1978), KOELN 22 103.

2. Cladoniafoliacea (Huds.) Willd. - Cladoniaceae Syn. Cladonia alcicornis (Lightf.) E. M. Fries. This terricolous, acidophytic to neutrophytic, Subcosmopolitan phyllolichen is

already known from all Macaronesian islands including the Cape Verdes (Santo Antao : Bouly de Lesdain 1935). Here it is reported from Sao Nicolau and Sao Tiago for the first time. According to the many collections available, which are not all cited below, it represents the characteristic species of the Cape Verdean subassociation of the soil-inhabiting Cladonietum foliaceae Klem. in the mountain belt.

San to Antao: Mossy ground, Lombo Cebide Vila south of Ribeira Grande, matt-forming, 1200 m, NW, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-9). - Mossy ground, Agua das Caldeiras, matt-forming, 1200 m, NE, pH 6,0; leg. W. Lobin (Xl /1979), FR CV-1342. - Mossy ground, north-east of Cova in the upper part of Ribeira do Paul, matt-forming, 1200 m, NE, pH 6,5; leg. 0. H. Rustan and C. Brochmann (1/1982), 0 (CV-68). Sao Nicolau: Mossy ground, northern slope of Monte Gordo, scattered, 900 m, NW, pH 6,5; leg. G . Follmann (IV/1974), KOELN 22 153. Sao Tiago: Open ground, associated with Cladonia rangiformis Hoffm., northern slope of Pico da Antonia, scattered, 1100 m, NE, pH 6,0; leg. G . Follmann (IV/l974), KOELN 22154.

3. Cladonia furcata (Huds.) Schrad. - Cladoniaceae Syn. Cladonia racemosa Hoffm.

This variable, terricolous, acidophytic, Cosmopolitan thamnolichen has previously been reported from all other Macaronesian archipelagos (Tavares 1952). For the Cape Verdes it is cited here for the first time. Whereas only one chemotype (with atranorin and fumarprotocetraric acid) has been found, in the collections mentioned below all main morphotypes can be observed : var. palamaea (Ach.) Nyl., var. pinnata (Floerke) Vain., var. racemosa (Hoffm.) Floerke. Obviously this ground lichen is of more sporadic occurrence in the cloud zone, especially in the dripping circles around epiphyte-covered shrubs.

Santo Antao : Gravelly ground, ridge between Cova and Ribeira do Paul, scattered, 1150 m, NE, pH 6,0; leg. P. Sunding (XI/1 976), 0 (CV-3). - Gravelly ground, Agua das Caldeiras, scattered, 1300 m, NE, pH 6,5; leg. W. Lobin (XI/1979), FR CV-1343, KOELN 22148. Sao Nicolau: Gravelly ground, summit of Monte Gordo and surroundings, scattered, 1000 m, NE, pH 6.5; leg. W. Lobin (XI/1980), FR CV-2231, KOELN 22 149. Sao Tiago: Gravelly ground, between Chao da Figueira and Pico da Malagueta, scattered, 900 m, NW, pH 6,5; leg. W. Lobin (X/1979), FR CV-461.

502 Journ . Hattori Bot. Lab. No. 61 1 9 8 6

4. Cladonia pocillum (Ach.) O. J. Rich. - Cladoniaceae This terricolous, basiphytic to neutrophytic, Subcosmopolitan phyllolichen has

already been collected on the Canary Islands and Madeira (Tavares 1952). It is new to the Cape Verde Islands, where it seems to be a sporadic companion species of the local subassociation of the soil-inhabiting Cladonietum foliaceae Klem. in the mountain belt.

Santo Antlio: Mossy ground, pass height between Porto Novo and Ribeira Grande, scarce, 1200 m, NW, pH 6,5; leg. K. Lewejohann (XII/1978), KOELN 22109. Sao Nicolau: Mossy ground, northern slope of Monte Gordo, scarce, 950-1100 m, NW, pH 6,5; leg. P. Sunding (XI/1976), KOELN 22 i56, o (CV-lO).

5. Cladonia rangiformis Hoffm. - Cladoniaceae Syn. Cladonia aetnensis Torn., Cladonia pungens (Ach.) Ach. In contrast to Cladonia furcata (Huds.) Schrad., this terricolous, acidophytic to

neutrophytic, Cosmopolitan thamnolichen has been reported from all Macaronesian archipelagos including the Cape Verde Islands (Santo Antao: Bouly de Lesdain 1935). First records are dealt with below for Sao Nicolau and Sao Tiago; only a few additional samples are cited from Santo Antao. Again the main morphotypes (var. muricata (Del.) Am., var. pungens (Ach.) Vain.) are present, whilst the spectrum of secondary compounds remains more or less constant: atranorin, ± fumarprotocetraric acid, rangiformic acid. According to our field notes, this is the most abundant larger fruticulose species of the Cladoniaceae in moister, partly protected, and shadowy niches with well drained lava soil in the mountain belt of the larger islands of the group.

San to Antlio: Gravelly soil, Lombo Cebide Vila south of Ribeira Grande, scattered, 1200 m, NW, pH 6,5; leg. P. Sunding (X/1972), 0 (CV-7). - Gravelly soi l, track from Montanha Forte de Leite to Ribeira do Paul, abundant, 1000-1100 m, NW, pH 6,0; leg. P. Sunding (XI/1976), 0 (CV-6). Gravelly soil, track from Cova to Ribeira do Paul, abundant, 930 m, NE, pH 6,5; leg. L. Borgen (1/1980), 0 (CV-4). Slio Nieolauy: Gravelly soil, northern slope of Monte Gordo, scattered, 950-1100 m, NW, pH 6,5 ; leg. P. Sunding (XI/1976), 0 (CV-8). - Gravelly soil, summit region of Monte Gordo, scattered, 1000 m, NE, pH 6,5; leg. W. Lobin (XI/2980), FR CV-2234, KOELN 22 155, Slio Tiago: Gravelly soil, northern slope of Pico da Antonia, abundant, 1100 m, NE, pH 6,0; leg. G. Follmann (IV/1974), KOELN 22 151.

6. Diploschistes scruposus (Schreb.) Norm. - Diploschistaceae Syn. Lecanora scruposa (Schreb.) Nyl. , Parmelia seruposa (Schreb.) Hepp, Psora scruposa (Schreb.) Hampe.

This generally saxicolous, occasionally terricolous, acidophytic, Cosmopolitan lepolichen is already known from the Azores, Canaries, and Madeira (Tavares 1952). It is new to the Cape Verde Islands, where it can be considered a relatively rare rockinhabitant of half-shadowy, dew-wetted boulders and cliffs in the mountain belt.

Fogo: Flawy lava rocks, outer crater edge above Monte Velho, scarce, 1800 m, NE, pH 6,5; leg. G. Follmann (I1I/2986), KOELN 30017. Slio Nieolau: Lapilli on rocky slope, summit sphere of Monte Gordo, scarce, 1200 m, NE, pH 6,5 ; leg. W. Lobin (XI/1980), FR CV-2233, KOELN 30 018.

7. Graphis scripta (L.) Ach. - Graphidaceae Syn. Graphis hieolor Fee, Graphis nematodes Stirt., Opegrapha scripta (L.) Ach.

G. FOLLMANN & B. MIEs: Lichen flora and vegetation of Cape Verde Islands. IV 503

This variable, corticolous, acidophytic, Cosmopolitan lepolichen has been reported previously from the Canary Islands (Tavares 1952). It is new to the Cape Verde Islands. The few thalli available are well developed; morphologically they correspond to var. limitata (Pers.) Ach. of negligible taxonomic value. It seems to he a rare epiphyte of deciduous trees in the mountain belt.

Sao Tiago: Smooth bark of Faidherbia albida (Del.) Chev., Cruz de Goto Bravo, Serra da Pico. da Antonia, scattered, 1000 m, pH 6,0; It''g. G. Follmann (IV/1974), KOELN 22167, KOELN 22168.

8. Leprocaulon microscopicum (ViII.) Gams - Leprariaceae Syn. Leprocaulon nanum (Ach.) Nyl., Stereocaulon microscopicum (ViII.) Frey, Stereocaulon quisquiliare (Leers.) Hoffm.

Occurring on all Macaronesian archipelagos, this saxicolous or terricolous, rarely corticolous, acidophytic to neutrophytic, Subcosmopolitan deuterolichen has already been recorded from Siio Nicolau and Sao Tiago (Tavares 1964). It is new to Santo Antao and Sao Vicente. In contrast to the other Macaronesian groups, on the Cape Verde Islands it is always rare and poorly developed. However, it shows a relatively complete spectrum of secondary products ("deficient phase VII" of Mackenzie-Lamb and Ward 1974, with atranorin, usnic acid, and zeorin).

Santo Antao: Sandy loam, north-west of Cova in the upper part of Ribeira do Paul, scarce, 1130 m, SE, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-2). - Loam accumulations, track from Montanha Forte de Leite to Ribeira do Paul, scarce, 1000-1100 m, NW, pH 6,5; leg. P. Sunding (XI/1 976), 0

(CV-I). Sao Vicente: Gravelly loam, north-eastern slope of Monte Verde near Jaime, scarce, 600 m, NE, pH 6,5; leg. G. Follmann (IV/1 974), KOELN 22147.

9. Leptogium lichenoides (L.) Zahlbr. - Collemataceae Syn. Leptogium atrocaeruleum (Schaer.) Mass., Leptogium lacerum (Raeusch.) S. F . Gray.

This generally corticolous or muscicolous, occasionally saxicolous, euryecious, Cosmopolitan gleolichen has been cited for Madeira by Tavares (1952). It is new to the Cape Verde Islands, where it is known only in sterile condition on periodically moist cliffs in the cloud zone.

San to Antao: Shadowy, mossy rocks, north-west of Cova in the upper part of Ribeira do Paul, scarce, 1130 m, NE, pH 6,5; leg. P. Sunding (X/1 972), 0 (CV-66).

10. Parmelia borreri (J. E. Smith) Turn. - Parmeliaceae Syn. Parmelia aleuriza Vain., Parmelia borrerioides Nyl., Parmelia insignata Stiz. This corticolous or saxicolous, acidophytic to neutrophytic, Subcosmopolitan

phyllolichen with Oceanic distribution tendencies is already known from the Canaries (0sthagen & Krog 1976) and Cape Verdes (Santo Antao: 0sthagen 1974). A first record is presented here for Siio Tiago. As usual, no apothecia could be found, but the characteristic compounds (atranorin, gyrophoric acid) are in accordance with tho&e of the type material. On both islands it seems to be a rare bark inhabitant of well lit sites in the mountain belt.

Sao Tiago: Rough bark of Faidherbia albida (Dei.) Chev., south-western slope of Pi co da Antonia, scarce, 950-1050 m, NW, pH 6,5; leg. P. Sunding (XI/1976), 0 (CV-19).

11. Parmotrema chinense (Osb.) Hale et Ahti - Parmeliaceae Syn. Parmelia perlata (Huds.) Ach., Parmelia trichotera Hue, Parmotrema perlatum (Huds.) Choisy.

504 Journ. Hattori Bot. Lab. No. 61 198 6

This corticolous or saxicolous, acidophytic, SUbcosmopolitan phyllolichen is listed for all Macaronesian archipelagos by Tavares (1952). However, according to a remark by Zahlbruckner (Sao Vicente: 1906), its existence on the Cape Verde Islands has been doubtful till today. It is confirmed here for the Cape Verdes, where it seems to be an occasional epilithophyte in the cloud zone. Morphologically and chemically our collections are indistinguishable from African, American, Canarian, and European ones.

Santo Antao : Smooth bark of Eucalyptus globulus Lab., upper hollow of Ribeira Grande, scattered, 900 m, SE, pH 6,5; leg. G. Follrnann (IV/1 974), KOELN 22 162. Sao Nicolau: Dry shrubs, between Cacha<;o and Monte Gordo, scattered, 800-920 m, NE, pH 6,0; leg. P. Sunding (XI/1976), o (CV-25). Sao Tiago : Rough bark of Faidherbia albida (Del.) Chev., south-western slope of Pico da Antonia, scattered, 900 rn, NW, pH 6,5; leg. G. Follmann (IV/1 974), KOELN 22 163. - Halfshadowy, volcanic outcrops, east of summit of Pico da Malagueta, scattered, 950 m, NE, 6,0; leg. P. Sunding (XI/ l 976), 0 (CV-26).

12. Peltigera rufescens (Weiss) Humb. - Peltigeraceae Syn. Peltigera canina (L.) Willd. var. rufescens (Weiss) Willd.

This terricolous, basiphytic to neutrophytic, Subcosmopolitan phyllolichen has already been reported from the Azores, Canaries, and Madeira (Tavares 1952). It is new to the Cape Verde Islands. Together with Peltigera canina (L.) Willd. (Santo Antao: Bouly des Lesdain 1935) it can be considered a relatively rare companion species of the Cape Verdean subassociation of the soil-inhabiting Cladonietumfoliaceae Klem. in the mountain belt.

Santo Antao: Mossy soil with Cladonia foliacea (Huds.) Willd., track from Montanha Forte de Leite to Ribeira do Paul, scattered, 1000-1100 m, NE, pH 6,5; leg. P. Sunding (XI/1 976), 0 (CV-38). Sao Tiago: Mossy ground, north-western slope of Pico da Malagueta, scattered, 950 m, NW, pH 6,5; leg. G. Follmann (IV/1974), KOELN 22166.

13. Phaeophyscia orbicularis (Neck.) Mob. - Physciaceae Syn. Physcia obscura (Humb.) Fuern., Physcia orbicularis (Neck.) Poetsch, Physcia virella (Ach.) Flag.

This generally corticolous, occasionally saxicolous, relatively euryecious, Subcosmopolitan phyllolichen has been noted from the Azores and Canaries by Tavares (1952). It is new to the Cape Verde Islands. The only specimen available is well developed and fertile, but for the time being it must be considered a rare epiphyte of deciduous trees in the mountain belt.

Siio Tiago : Rough bark of Faidherbia albida (Del.) Chev., south-western slope of Pieo da Antonia, scattered, 950-1050 m, NW, pH 6,5; leg. P. Sunding (XI/1976), 0 (CV-71).

14. Physcia tenella (Scop.) de Cand. - Physciaceae Syn. Physcia hispida (Schreb.) Frege var. tenella (Scop.) Wats., Physcia marina (Nyl.) Lynge, Physcia subobscura (Nyl.) Nyl.

This corticolous or muscicolous, more rarely saxicolous, sUbacidophytic to subbasiphytic, Subcosmopolitan phyllolichen has been reported from the Azores by Tavares (1952) and from the Canary Islands by Sl'mchez-Pinto et al. (1983). It is new to the Cape Verde Islands. The sterile collection mentioned below corresponds to the


corticolous ecotype (var. tenel/a). Like the preceding, it can be considered a rare epiphyte of deciduous shrubs and trees in the mountain belt.

Slio Nicolau: Dead twigs, north-east of Monte Deserto, scarce, 800 m, NE, pH 6,0; leg. P. Sunding (XI/1976), 0 (CV-70).

15. Physcia tribacia (Ach.) Nyl. - Physciaceae Syn. Physcia caesitia Nyl. , Physcia erosa (Borr.) von Zwackh. This corticolous or saxicolous, subneutrophytic to subbasiphytic, Subcosmo

politan phyllolichen has been cited for the Canaries by Tavares (1952) and for Madeira by Arvidsson and Wall (1985). It is new to the Cape Verde Islands. Hitherto only a small but well developed and fertile sample is available. It seems to be a rare lithophyte in the mountain belt.

Santo Antlio : Shadowy rocks, north-east of Cova in the upper part of Ribeira do Paul, scattered, 1150 m, NE, pH 6,5; leg. P. Sunding (XI/J976), 0 (CV-75).

16. Pseudoparmelia caperata (L.) Hale - Parmeliaceae Syn. Parmelia caperata (L.) Ach., Parmelia euplecta Stirt., Parmelia subglauca Nyl. This generally corticolous, exceptionally saxicolous, acidophytic, Suboceanic,

Subcosmopolitan phyllolichen is known from all Macaronesian archipelagos (Tavares 1952), but has hitherto been recorded only once for the Cape Verde Islands (Sao Nicolau: Cardoso 1915). It is new to Santo Antao but must be regarded a rare lithophyte in the cloud zone. Nevertheless, the sterile specimen cited below is well developed and holds the complete set of secondary products (atranorin, caperatic acid, protocetraric acid, usnic acid).

Santo Antlio: Half-shadowy, volcanic outcrops, ridge between Cova and Ribeira do Paul, scarce, 1150 m, NE, pH 6,0 ; leg. P. i unding (XI/1976), 0 (CV-24).

TROPICAL GEOELEMENTS

17. Heterodermia isidiophora (Nyl.) Awas. - Physciaceae Syn. Anaptychia isidiophora (Nyl). Vain., Anaptychia speciosa (Wulf.) Mass. f. isidiophora (Nyl.) Zahlbr. This corticolous or saxicolous, acidophytic, Pantropical phyllolichen has been

reported from the Cape Verde Islands by Bouly de Lesdain (Fogo: 1935), but is apparently lacking on the other Macaronesian archipelagos. Although the record mentioned above is omitted in Kurokawa's monograph and supplement (1962, 1973), it is confirmed here for the Cape Verdes. To judge from our field observations, this often luxuriantly developed lithophyte represents a characteristic species of the Cape Verdean subassociation of the rock-inhabiting Parmotremetum tinctorum Follm. in the mountain belt.

Fogo: Open lava rocks, Ribeira Rocinha between Monte Velho and Ponta Gil, frequent, 900 m, NE, pH 6,5; leg. G. Follmann (111/1986), KOELN 22179. Santo Antlio: Lava outcrops, Monte Joana, frequent, 850-900 m, NW, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-13). - Lava outcrops, northeast of Cova in the upper part of Ribeira do Paul, numerous, 1030 m, NE, pH 6,0; leg. L. Borgen (1/1980),0 (CV-14). Slio Nicolau: Detritus-covered rocks, east of Tope de Matim, scattered, 600 m, NE, pH 6,0; leg. P. Sunding (XI/1976), 0 (CV-IS). Slio Tiago: Smooth bark of Acacia ni/otica (L.)


Del. and Lantana camara L. , Monte Campanario, scattered, 1000 m, NE, pH 6,5; leg. P. Sunding (XI/1972), 0 (CV-17). - Rough bark of Faidherbia albida (Del.) Chev., south-western slope of Pico da Antonia, scattered, 950-1050 m, NW, pH 6,0; leg. P. Sunding (XI/1976, 0 (CV-16).

18. Heterodermia japonica (Sato) Swinsc. et Krog - Physciaceae Syn. Anaptychia chilensis Kur. var. austroafricana Kur., Anaptychia dendritica (Pers.) Vain. var. japonica Sato, Anaptychia japonica (Sa to) Kur. This relatively substrate-vague, acidophytic, Paleotropical phyllolichen has been

collected on Madeira by Arvidsson and Wall (1985) and on the Canries by 0sthagen and Krog (1976). It is new to the Cape Verde Islands. In contrast to Heterodermia isidiophora (Nyl.) Awas., it seems to be a rare lithophyte of shadowy, mossy, volcanic rocks in the cloud zone.

Santo Antao : Shadowy, mossy cliffs, Lombo Cebide Vila south of Riberia Grande, scarce, 1200 m, SW, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-77).

19. Leptogiumfurfuraceum (Harm.) Sierk. - Collemataceae Syn. Leptogium hildenbrandii (Gar.) Nyl. f. furfuraceum Harm.

This generally corticolous, sometimes muscicolous, acidophytic, disjunct Subtropical gleolichen has been recorded for Madeira (Tavares 1952). Hitherto found only in steril condition, it is new to the Cape Verde Islands and can be considered one of the rare epibryophytes in the mountain belt.

Santo Antao: Shadowy, mossy rocks, Lombo Cebide Vila south of Ribeira Grande, scarce, 1100 m, NE, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-64). Sao Vicente : Shadowy, mossy rocks, plateau of Monte Verde, scarce, 700 m, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-65).

20. Parmotrema reticulatum (Taly.) Choisy - Parmeliaceae Syn. Parmelia cetrata Hue, Parmelia perforata (1. E. Smith et Sow.) S. F. Gray, Parmelia reticulata Tay!.

This corticolous or saxicolous, acidophytic, Pantropical phyllolichen is known from all Macaronesian islands (Tavares 1952). Bouly de Lesdain (1935) cited it for Santo Antiio and Follmann and Sanchez-Pinto (1986) found it on Sao Tiago and Sao Vicente. Here it is reported from Fogo and Sao Nicolau for the first time and additional findings are noted from Santo Antao. In contrast to other Macaronesian biotopes, on the Cape Verde Islands the epilithic ecotype clearly predominates. It represents one of the most aggressive and remarkable characteristic species of the Cape Verdean subassociation of the rock-colonizing Parmotremetum tinctorum Follm. in the mountain belt.

Fogo: Volcanic rocks, path from Monte Velho to Mosteiros, l100m, NE, pH 6,5; leg. G. Follmann (111/1986), KOELN 30 002. Santo Antao : Volcanic rocks, north-western slope of Monte Joana, abundant, 850-900 m, NW, pH 6,0; leg. P. Sunding (X/1 972), 0 (CV-31). - Volcanic rocks, upper part of Ribeira das Pedras, scattered, 750-900 m, NE, pH 6,5; leg. P. Sunding (XI/1976), 0

(CV-32). - Volcanic rocks, north-east of Cova in the upper part of Ribeira do Paul, abundant, 1030 rn, NE, pH 6,0; leg. L. Borgen (1/1980), 0 (CV-40). Sao Nicolau: Dry shrubs, between Cacha~o and Monte Gordo, scattered, 900 m, NE, pH 6,5; leg. P. Sunding (Xlj2976), 0 (CV-29). - Volcanic rocks, northern slope of Monte Deserto facing Ribeira da Fragata, scattered, 900 m, NE, pH 6,0; leg. P. Sunding (XljI976), 0 (CV-27). - Volcanic rocks, east of Tope de Matim, abundant, 600 m, NW, pH 6,0; leg. P. Sunding (XI/1976), 0 (CV-89).

G. FOLLMANN & B. MIEs: Lichen flora and vegetation of Cape Verde Islands. IV 507

21. Ramalina peruviana Ach. - Ramalinaceae This mostly corticolous, sometimes saxicolous or terricolous, acidophytic, Pan

tropical thamnolichen is new for the Macaronesian floral region. Possibly, collections from the Cape Verde Islands have been confused with the externally similar Ramalina dendriscoides Nyl. (San to Antao, Sao Tiago: Bouly de Lesdain 1935), which differs in having most soralia situated apically on short lateral branchlets and in containing salazinic acid, and which could not be refound till today. Appearing in both morphotypes (var. pollinariaeformis Vain., var. tenuescens Vain.), this epiphyte is a frequent companion species of the Cape Verdean subassociation of the Teloschistetum flavicantis Follm. of the open shrub formation in the cloud zone.

Sal: Dry shrubs, summit of Monte Grande, scattered, 400 m, NW, pH 6,0; leg. G. Follmann (111/1986), KOELN 40 003. Siio Nicolau: Dry twigs, eastern and southern slopes of Alto da Joaquina, scattered, 550-600 m, NE, pH 6,5; leg. P. Sunding (XI/1976), 0 (CV-40). - Gravelly soil, between Cachar;o and Monte Gordo, scattered, 800-920 m, NE, pH 6,5 ; leg. P. Sunding (XI/1976), 0 (CV-41). - Dry shrubs, eastern surroundings of Monte Gordo, scattered, 1000 m, NE, pH 6,5; leg. W. Lobin (XII/1978), FR CV-1261, KOELN 22105. Siio Tiago : Dry shrubs, west of summit of Pieo da Malagueta, scattered, 860 m, NW, pH 6,5; leg. P. Sunding (XI/1976), 0 (CV-42), - Wooden post, Ribeira Janela between Rui Vaz and Sao Jorge, abundant, 820 m, N, pH 7,0; leg. 0. H . Rustan (1/ 1980),0 (CV-43). Siio Vicente: Dry twigs, plateau of Monte Verde, abundant, 700 m, NE, pH 6,5; leg. P. Sunding (X/1972), 0 (CV-46). - Smooth leaves of Furcraea gigantea Vent., near top of Monte Verde, scattered, 700 m, NW, pH 6,0; leg. P. Sunding (XI/1976), 0 (CV-44). - Volcanic outcrops, north-eastern slope of Monte Verde, frequent , 590-660 m, NE, pH 6,5 ; leg. L. Borgen (1/1980) , 0

(CV-45).

22. Roccella africana Vain. - Roccellaceae This saxicolous, acidophytic, halophytic, Paleotropical thamnolichen with a

Namibian distribution centre has already been recorded for the Azores (Tavares 1952). It is new to the Cape Verde Islands, where it can be considered a characteristic species of the local subassociation of the epilithic Roccelletum canariensis Follm. in the lower mountain belt.

Boa Vista: Volcanic rocks, northern slope of Monte Rochinha north of Sal-Rei, scattered, 50-90 m, NW, pH 6,5; leg. L. Borgen (1/1980), 0 (CV-84). Sal: Volcanic rocks, eastern slope of Monte Grande, abundant, 200-300 m, NE, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-85). - Volcanic rocks, Monte Rocha da Salina, abundant, 300 m, NW, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-86). Siio Nicolau: Lava face, north-eastern slope of Pico Ventoso, 700 m, NE, pH 6,0; leg. G. Follmann (111/ 1986), KOELN 30004.

23. Teloschistes villosus (Ach.) Norm. - Teloschistaceae Syn. Anaptychia villosa (Ach.) Boist., Physcia villosa (Ach.) Duby.

This generally corticolous, occasionally saxicolous, acidophytic to neutrophytic, disjunct Subtropical thamnolichen with a West Mediterranean distribution centre has hitherto been noted from the Canary Islands (Tavares 1952). It is new to the Cape Verde Islands. All samples seen (about 50) represent a fine-lobed morphotype, and only a single fertile thallus has been observed. On Sal and Sao Nicolau this epiphyte grows together with Tornabea atlantica (Ach.) 0sth. in the Cape Verdean subassociation of the Teloschistetum flavicantis Follm. in the mountain belt.


FIG. 2. Roccelletum canariensis Follm. with Gorgadesia mira Tav. (fruticulose) and Lecanora sulphurella Hepp (crustaceous) on volcanic rocks in the lower mountain belt of SaD Vicente (1986).

Sal: Dry shrubs, summit of Monte Grande, 400 m, NW, pH 6,0; leg. G. Follmann (111/1986), KOElN 30 005. Sao Nicolau: Dry shrubs, north-eastern slope of Monte Agua de Jose, 650 m, NE, pH 6,5 ; leg. G . Follmann (III/1986), KOELN 30002.

24. Usnea baylei (Stirt.) Zahlbr. - Usneaceae Syn. Usnea formosa (Stirt.) Zahlbr., Usnea implicita (Sti rt.) Zahlbr., Usnea insignis Mot. This generally corticolous, exceptionally saxicolous, acidophytic, Pantropical

thamnolichen has been reported from the Cape Verde Islands and Madeira as Usnea implicita (Stirt.) Zahlbr. (Tavares 1952). Unfortunately, no exact collection data are indicated for the first island group (Bouly de Lesdain 1935) ; therefore it is new to Sao Nicolau and Sao Vicente. At present it can be taken for a relatively rare lithophyte of the lower cloud zone.

sao Nicolau: Stone wall, south-east of Alto das Caba<;as, scattered, 550 m, S, pH 6,5; leg. 0. H. Rustan & C. Brochmann (II/1982), 0 (CV-62), saG Vicente: Volcanic outcrops, north-western slope of Monte Verde, scarce, 550 m, NW, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-61).


MEDITERRANEAN GEQELEMENTS

25. Dirina massiliensis Our. et Mont. - Roccellaceae Syn. DirinaJallax de Not., Dirina patronii Bag\., Dirina repanda E. M. Fries. Since Crombie's revision of the lichens of the Challenger Expedition to the

Midatlantic Islands (1877), Dirina repanda E. M. Fries has been cited without exact collection site for the Cape Verde Islands (Bouly de Lesdain 1935, Tavares 1952), but Tehler denied the existence of the saxicolous, acidophytic to basiphytic, halophytic, Atlantic-Mediterranean lepolichen in the Macaronesian floral region. Meanwhile, Herm'mdez-Padron et al. (1986) reported it from the Canary Islands, and now a well developed population has been discovered on the Cape Verde Islands. Actually, it can be considered a rare companion species of the Cape Verdean subassociation of the epilithic Roccelletum canariensis Follm. in the lower mountain belt.

Sal: Volcanic boulder, among fruticulose Roccellaceae, western base of Monte Grande, scarce, 100 rn, NW, pH 6,5; leg. G. Follrnann (III/1986), KOELN 30001. Sao Nicolau: Volcanic rock face, together with fruticulose Roccellaceae, heights south-east of Cacha9a, scattered, 600 rn, NE, pH 6,5; leg. G. Follrnann (1lI/1986), KOELN 30 013.

26. Heppia turgida (Ach.) Nyl. - Heppiaceae Syn. Endocarpon turgidum Ach., Gloeoheppia turgida (Ach.) Gyeln.

This normally terricolous, exceptionally saxicolous, basiphytic, Atlantic-Mediterranean gleolichen has recently been cited for the Canary Islands by Sanchez-Pinto et al. (1983) and Swinscow (1982). It is new to the Cape Verde Islands, where it seems to be a rare pioneer lichen on bare mineral soil in the mountain belt.

San to Antao: Sandy soil, terraces in the upper part of Ribeira Grande, scarce, 600 rn, NE, pH 7,0; L. Sanchez-Pinto (VIII/1978), KOELN 22 142.

27. Peltigera hymenina (Ach.) Del. - Peltigeraceae Syn. Peltigera polydactyla (Neck.) Hoffrn. var. crassoides Gyeln., Peltigera polydactyla (Neck.) Hoffrn. f. hymenina (Ach.) Flot., Peltigera rufescens (Weiss) Hurnb. var. hymenina (Ach.) Hepp .

This primarily terricolous, also muscicolous or saxicolous, euryecic, AtlanticMediterranean phyllolichen is new to the Macaronesian floral region, but has probably been listed before under Peltigera polydactyla (Neck.) Hoff'm. (Tavares 1952). The only gathering available for the moment shows well developed but sterile thalli. Apparently it is a rare companion species of the Cape Verdean subassociation of the soil-inhabiting Cladonietum foliaceae Klem. in the mountain belt.

Santo Antao: Gravelly soil with Cladonia rangiformis Hoffrn., north-west of Cova in the upper part of Ribeira do Paul, scattered, 1240 rn, NE, pH 6,5; leg. P. Sunding (X/1972), 0 (CV-79).

28. Pertusaria texana Muell.-Arg. - Pertusariaceae Syn. Pertusaria disticha Erichs.

This normally corticolous, occasionally saxicolous, acidophytic, South AtlanticNorth American lepolichen is new to Macaronesia. Our sole sample corresponds in all essential morphological, anatomical, chemical, and ecological characters to the detailed description given by Dibben (1980) and has been compared with North

510 Journ. Hattori Bot. Lab. No. 61 1 9 8 6

American collections. It can be considered a rare epiphyte of solitary, deciduous trees in the mountain belt.

sao Tiago: Smooth bark of Faidherbia albida (Del.) Chev., north-eastern slope of Pi co da Antonia, scattered, 1100 m, N, pH 6,0; leg. G. Follmann (lV/1974), KOELN 22171.

29. Psora tabacina (Ram.) de Cand. - Psoraceae Syn. Biatora tabacina (Ram.) E. M. Fries, Lecanora tabacina (Ram.) Steud., Lecidea tabacina (Ram.) Schaer.

This terricolous, basiphytic to neutrophytic, Macaronesian-Mediterranean phyllolichen has already been recorded for the Canaries and Madeira (Tavares 1952). It is new to the Cape Verde Islands, colonizing there smaller, partly protected, horizontal loam accumulations between lava boulders in the cloud zone.

sao Nicolau: Sandy loam, north-eastern slope of Pico Ventoso, spot-l ike, 800 m, NW, pH 7,0; leg. G. Follmann (III/1986), KOELN 30019.

30. Ramalina duriaei (de Not.) Bagl. - Ramalinaceae Syn. Ramalina evernioides auct., Ramalina pollinaria (Westr.) Ach . var. duriaei de Not., Ramalina pusilla le Prev. f. dilacerala Mass. This corticolous or saxicolous, acidophytic, Macaronesian-Mediterranean phyl

lolichen is known from all Macaronesian archipelagos including the Cape Verde Islands (Brava, Sao Vicente: Follmann & Sanchez-Pinto 1986). First records are dealt with here for Sal, Sao Nicolau, and Sao Tiago; moreover, additional ones are given for Sao Vicente. Most of the samples studied belong to the epilithic ecotype and tend morphologically to f. divisa Wern. and f. fistulosa Wern. of negligible taxonomic value. At none of the locations cited below this lithophyte reaches a higher population density. It represents a sporadic companion species of the Cape Verdean rock-inhabiting Ramalinetum maderensis Follm. et Mies in the lower mountain belt.

Sal: Older twigs of Euphorbia tuckeyana Steud., eastern slope of Monte Grande, scattered, 300 m, NE, pH 6,0 ; leg. P. Sunding (X/1972), 0 (CV-71). sao Nicolau : Volcanic tuff, south-east of Cachaya in the upper part of Ribeira das Patas, scattered, 600 m, NW, pH 6,5 ; leg. L. Borgen (1/1 980), 0 (CV-73). sao Tiago : Volcanic rocks, west of summit of Pico da Malagueta, scattered, 950 m, NW, pH 6,0; leg. L. Borgen (1/1980), 0 (CV-72). sao Vicente: Volcanic tuff, northern slope of Tope da Caixa, scattered, 440 m, NW, pH 6,5; leg. P. Sunding (X/1972), 0 (CV-75). - Volcanic rocks, Gao Alto, scattered, 350-400 m, NE, pH 6,0; leg. P. Sunding (XI/1976), 0 (CV-74). - Volcanic rocks, Topinho do Feijoal, scattered, 350 m, NW, pH 6,5; leg. P. Sunding (XJ/1976), 0 (CV-76).

31. Ramalina maci/ormis (Del.) Bory - Ramalinaceae Syn. Desmazieria maci/ormis (Del.) Follm., Niebla maciformis (Del.) Bowl. et Rund., Ramalina evernioides Nyl.

This saxicolous, acidophytic, Atlantic-Mediterranean phyllolichen has its distribution centre in North Africa but occurs also on the Canary Islands and on Madeira (Krog & Osthagen 1980). It is new to the Cape Verde Islands. Morphologically, all Cape Verdean gatherings correspond to f. laciniata Wern., a growth form without taxonomic significance. On the other side, but in accordance with the situation on the Canary Islands, morphologically indistinguishable chemotypes with bourgeanic, norstictic, and usnic acids (I) and with bourgeanic, salazinic, and usnic acids (11) ap-


FIG. 3. Ramalinetum maderensis Follm. et Mies (left-hand) with Ramalina maderensis Mot. overgrowing Lecanora sulphurella Hepp and Roccelletum canariensis Follm. (righthand) with Roccella vicentina (Vain.) Follm. on volcanic boulder in the lower mountain belt of Sao Vicente (1974).

pear on the Cape Verde Islands. Like the preceding Ramalina duriaei (de Not.) Bag!., this lithophyte is an occasional companion species of the Cape Verdean rock-inhabiting Ramalinetum maderensis Follm. et Mies in the lower mountain belt.

Santo Antao: Volcanic rocks, Lombo Cebide YHa south of Ribiera Grande, scattered, 1100 m, NE, pH 6,0; leg. P. Sunding (XI/l976), 0 (CV-36). - Volcanic rocks, track from Montanha Forte de Leite to Ribeira do Paul, scattered, 1000- 1100 m, NE, pH 6,5; leg. P. Sunding (XljI976), 0 (CV-34). Sao Nicolau: Volcanic tuft', north-eastern slope of Pico da Cachar;a, scattered, 1000 m, NE, pH 6,5; leg. G. Follmann (IV/l974), KOELN 22164.

512 Journ. Hattori Bot. Lab. No. 61

32. Ramalina subgeniculata Nyl. - Ramalinaceae Syn. Ramalina aulota Stirt.

1 9 8 6

This corticolous or saxicolous, acidophytic, Macaronesian-Mediterranean thamnolichen has been reported from all Macaronesian archipelagos, the record for the Cape Verde Islands being based on a collection from Sao Vicente (Bouly de Lcsdain \935). The fistulose, divaricatic acid and usnic acid containing epilithophyte is confirmed here for the Cape Verdes and noted from Santo AnUio, Sao Nicolau, and Sao Tiago for the first time. Whilst Krog and 0sthagen (1980) found only the epiphytic ecotype on the Canaries, on the Cape Verdes epiphytic and epilithic ones occur. And whereas Ramalina subgeniculata Nyl. is a widespread, characteristic species of the bark-inhabiting Ramalinetum subgeniculatae Klem. on the Canary Islands, it is a rare companion species of various other lichen associations in the lower mountain belt of the Cape Verde Islands.

Santa Antao: Volcanic rocks, upper part of Ribeira Grande, scarce, 600 m, NE, pH 6,0; leg. L. Sanchez-Pinto (VIII/1978), KOELN 22170. Sao Nicolau: Dry twigs, east of Tope de Matim, scattered, 600 m, NE, pH 6,0; leg. P. Sunding (XI/1976), 0 (CV-49). Sao Tiago: Smooth stems of Ceratonia siliqua L., upper part of Ribeira da Janela between Rui Vaz and Sao Jorge, abundant, 670 m, NW, pH 6,0; leg. L. Borgen (1/1980), 0 (CV-51). - Dead twigs of Eucalyptus globulus Lab., upper part of Ribeira da Janela between Rui Vaz and Sao Jorge, abundant, 680 m, NE, pH 6,5; leg. L. Borgen (1/1980), 0 (CV-78). - Old wood of Eucalyptus globulus Lab., upper part of Ribeira da Janela between Rui Vaz and Sao Jorge, scattered, 600 m, NW, pH 6,5; leg. 0. H. Rustan (1/1980),0 (CV-50). Sao Vicente: Older shoots of Euphorbia tuckeyana Steud., north-western slope of Monte Verde, scattered, 520 m, NW, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-53). - Volcanic rocks, northwestern slope on Monte Verde, scarce, 460 m, NW, pH 6,5; leg. P. Sund ing (XI/1976), 0 (CV-54).

33. Roccella allorgei des Abb. - Roccellaceae This saxicolous, acidophytic, halophytic, Macaronesian-Mediterranean tham

nolichen has been found on the Azores (Tavares 1952) and the Canary Islands (Hernandez-Padron et al. 1986). It is new to the Cape Verde Islands, where it can be considered a rare companion species of the Cape Verdean subassociation of the epilithic Roccelletum canariensis Follm. in the lower mountain belt.

Boa Vista : Volcanic rocks, northern slope of Monte Rochinha north of Sal-Rei, scattered, 50-90 m, pH 6,5; leg. L. Borgen (1/1980), 0 (CV-87), Sal: Volcanic rocks, north-western slope of Monte Grande, scattered, 300 rn, NW, pH 6,0; leg. G. Follmann (III/1986), KOELN 30006. Sao Nicolau : Volcanic rocks, upper part of Ribeira das Patas south of Cacha<;a, scattered, 600 m, NE, pH 6,0; leg. L. Borgen (1/1980), 0 (CV-88).

34. Squamarina cartilaginea (With.) James - Squamarinaceae Syn. Lecanora benacensis (Mass.) Jatta , Lecanora crassa (Huds.) Ach., Squamarina crassa (Huds.) Poelt. This variable, terricolous, basiphytic to neutrophytic, Macaronesian-Mediter

ranean phyllolichen is known from the Canary Islands and Madeira (Tavare!> 1952) and the Cape Verde Islands (Sao Tiago: Follmann & Sanchez-Pinto 1986). A first record is mentioned here for Santo Antao. In contrast to the Canary Islands, where it is a characteristic species of the terricolous Diploschistetum albescentis Klem., it seems to be a relatively rare pioneer lichen on bare mineral soil in the mountain belt


FIG. 4. Teloschistetum fiavicantis Follm. with Teloschistes fia vicans (Swartz) Norm. and Tornabea atlantica (Ach.) 0sth. on Euphorbia tuckeyana Steud. on lava slope in the middle mountain belt of Siio Nicolau (\986).

of the Cape Verde Islands. San to Antiio : Sandy loam, Lombo Cebide Vila south of Ribeira Grande, scarce, 1100 m, NW,

pH 7,0; leg. P. Sunding (X/1972).

35. Tornabea atlantica (Ach.) 0sth. - Physciaceae Syn. Anaptychia intricata (Desf.) Mass., Tornabenia intricata (Desf.) Trev., Tornabeniopsis atlantica (Ach.) Follm. This corticolous or saxicolous, acidophytic, Macaronesian-Mediterranean tham

nolichen has been reported from the Canary Islands and Cape Verde Islands (Tavares 1953). The record for the latter island group is based on Anaptychia intricata (Desf.) Mass. var. denudata Bouly de Lesd. (SflO Tiago: Bouly de Lesdain 1935), an unstable morphotype which does not deserve taxonomic recognition (Kurokawa 1962). It is documented here for Sal, Santo Antao, and Sao Nicolau for the first time. All ecotypes and morphotypes occur among the collections cited below: applanate or


cylindrical and epilithic or epiphytic. It can be considered a substrate-vague companion species of the Cape Verdean subassociation of the hygrophytic Teloschistetum flavicantis Follm. in the cloud zone.

Sal: Older twigs of Euphorbia tuckeyana Steud., eastern slope of Monte Grande, scarce, 300 rn, NW, pH 7,0; leg. P. Sunding (X/1972), 0 (CV-ll). San to Antao: Side branches of Cupressus macrocarpa Hartw., pass height between Porto Novo and Ribeira Grande, scarce, 1200 rn, NW, pH 6,0 ; leg. G. Follrnann (IV/1 974), KOELN 22157. - Dry shrubs, upper part of Ribeira da Torre, scarce, 800 rn, NE, pH 6,5 ; leg. L. Sanchez-Pinto (VIII/1978), KOELN 22 166. Sao Nicolau: Lava outcrops, eastern and southern slopes of Alto da Joaquina, scarce, 550-600 rn, NE, pH 6,5; leg. P. Sunding (Xr/1976), 0 (CV-12). Sao Tiago : Volcanic rocks, south-western slope of Pi co da Antonia, scattered, 900 rn, NE, pH 6,0; leg. G. Follrnann (IV /1974), KOELN 22 172.

36. Xanthoparmelia tinctina (Mah. et Gill.) Hale - Parmeliaceae Syn. Parmelia rosea Gyeln" Parmelia tinctina Mah. et Gill. , Parmelia tokajensis Gyeln. This saxicolous, acidophytic, Atlantic-Mediterranean phyllolichen has recently

been found on the Canary Islands by Sanchez-Pinto et al. (1983). It is new to the Cape Verde Islands. At present it can be taken for an infrequent companion species of the Cape Verdean subassociation of the epilithic Parmotremetum tinctorum Follm. in the mountain belt.

Fogo: Lapilli-covered ground, ascent to Pico Novo, scarce, 2100 rn, NW, pH 6,0; leg. W. Lobin (X/l981), FR CV-1958, KOELN 22 173. Santo Antao: Volcanic rocks, Lornbo Cebide Vila south of Ribeira Grande, scarce, 1200 rn, NW, pH 6,5; leg. P. Sunding (X/1972), 0 (CV-69).

MACARONESIAN GEOELEMENTS

37. Gorgadesia mira Tav. - Roccellaceae This saxicolous, acidophytic, halophytic thamnolichen is one of the few local

endemics among the lichenophytes of the Cape Verde Islands. Hitherto it has been reported from Brava and Sao Tiago by Tavares (\964) and from Sao Vicente by Follmann (1974). First records are noted here for Sal, Santo Antao, and Sao Nicolau. On half-shadowy, perpendicular, volcanic rock faces it sometimes dominates the Cape Verdean subassociation of the hygrohalophytic Roccelletum canariensis Follm. in the lower mountain belt (differential species).

Sal: Volcanic rocks, north-western slope of Monte Grande, abundant, 300 rn, pH 6,0; leg. G. Fol\rnann (IV/1974), KOELN 22 174. Santo Antao: Volcanic rocks, coastal hills inland of Ponta do Sol, scattered, 200 rn, N, pH 6,0 ; leg. G. Fol\rnann (IV/l974), KOELN 22175. Sao Nicolau: Volcanic rocks, coastal cliffs near Estancia Braz, scattered, 400 rn, pH 6,5; leg. G. Follrnann (IV/1974), KOELN

22 176. Sao Vicente: Volcanic rocks, north-eastern slope of Tope da Suzana, scattered, 240 rn, pH 6,5; leg. P. Sunding (XI/2972), 0 (CV-80). - Volcanic rocks, northern slope of Monte Verde, abundant, 600 rn, NW, pH 6,0; leg. G. Follrnann (III/1 986), KOELN 40 007.

38. Ramalina bourgaeana Mont. - Ramalinaceae Syn. Desmazieria bourgaeana (Mont.) Follrn., Niebla bourgaeana (Mont.) Rund. et Bowl., Ramalina vulcania (Mont.) Nyl. This saxicolous, acidophytic, subhalophytic phyllolichen is known from Madeira,

the Canary Islands, and the Cape Verde Islands (Tavares 1952). Bouly de Lesdain (1935) cited it for Santo Antao and Follmann and Sanchez-Pinto (1986) for Sao


Vicente; it is new to Sal and Sao Nicolau. In contrast to the Canary Islands. on the Cape Verde Islands the deap-lacerate morphotype (f. delicata J. Stein.) clearly predominates. On the other side, in addition to chemotype I with bourgeanic acid, salazinic acid, triterpenoids, and usnic acid (Follmann & Sanchez-Pinto 1986), now chemotype 11 without salazinic acid has also been observed . Whilst it acts on the Canary Islands as the most characteristic and competitive species of the Ramalinetum bourgaeanae Follm. on maritime rocks, it is a sporadic companion species of the hygrohalophytic Ramalinetum maderensis Follm. et Mies which replaces the latter on the Cape Verde Islands.

Sal: Volcanic rocks, north-western slope of Monte Grande, scarce, 300 m, pH 6,0; leg. G . Follmann (IV/1974), KOELN 22 177. San to Antao : Volcanic rocks, Lombo Cebide Vila south of Ribeira Grande, scarce, 1100 m, NW, pH 6,5; leg. P. Sunding (X/1972), 0 (CV-8I). - Volcanic rocks, surroundings of Cova in the upper part of R ibeira do Paul, scarce, 1150 m, pH 6,0; leg. P. Sunding (XI/1976), 0 (CV-82). Sao Nicolau: Volcanic rocks, Cab~a de Vaca, scarce, 400 m, NW, pH 6,0; leg. G. Follmann (IV/1974), KOELN 22 178. Sao Vicente: Volcanic rocks, southern slope of Monte Vigia facing Mindelo, scarce, 280 m, pH 6,5; leg. P. Sunding (XI/1976), 0 (CV-83).

39. Ramalina maderensis Mot. - Ramalinaceae This saxicolous, acidophytic, subhalophytic thamnolichen has been collected on

Madeira, the Canaries, and the Cape Verdes (Sao Vicente : Motyka 1960). It is new to Boa Vista, Sal, Santo Antao, and Sao Nicolau. Like on the Canary Islands (Krog & 0sthagen 1980), both chemotypes (1 with lecanoric acid and usnic acid, 11 with 4-0-demethylbarbatic acid and usnic acid) occur on the Cape Verde Islands in more or less equal percentage. In view of the superficial similarity with some rigid morphotypes of Ramalina siliquosa (Huds.) A. L. Smith which does not reach the Cape Verde Islands, two subspecific taxa (var. cuspidata (Ach.) Magn. f. crassa (Nyl.) Tav., var. erythrocarpa (Mont.) Tav.) proposed and listed by Tavares (1952) can probably be placed here. In the Barlavento Group it is one of the most common and striking pioneer lichens on exposed volcanic rocks above the Roccellaceae belt. For that very reason it has been selected as naming species of the Cape Verdean Ramalinetum maderensis Follm. et Mies which will be described in detail elsewhere.

Boa Vista : Volcanic rocks, Morro Preto near Punta do Porto Ferreira, scattered, 100 m, NE, pH 6,5; leg. G. Follmann (IV/1974), KOELN 22 158. Sal : Volcanic rocks, top of Monte Rocha da Salina, abundant, 300 m, NW, pH 6,5; leg. P. Sunding (X/1972), 0 (CV-33). - Coastal rocks, Punta da Cagada near Bahia Calhetinha, abundant, 50 m, NE, pH 6,5; leg. G . Follmann (lV/1974), KOELN 22 159. - Volcanic rocks, summit region of Monte Grande, abundant, 400 m, NW, pH 6,0; leg. G . Follmann (III/1986), KOELN 30008. Santo Antao: Volcanic rocks, Punta da Tumba in front of I1h6u Lombo de Boi, abundant, 150 m, NE, pH 6,0 ; leg. G. Follmann (IV/1974), KOELN 22 160. Sao Nicolau : Volcanic rocks, southern and eastern slopes of Alto da Joaquina, abundant, 550-600 m, NE, pH 6,5; leg. P. Sunding (XI/1976), 0 (CV-20). - Volcanic rocks, south-east of Tope de Matim, abundant, 770 m, S, pH 6,0; leg. 0. H . Rustan (1/1980), 0 (CV-21). Sao Vicente : Volcanic rocks, north-western slope of Monte Verde, abundant, 350 m, NW, pH 6,0 ; leg. G. Follmann (IV/1974), KOELN 22 161. - Volcanic rocks, surroundings of Gao Alto, abundant, 350-400 m, N, pH 6,0; leg. P. Sunding (XI/1967), 0 (CV-22). - Volcanic rocks, southern slope of Monte Vigia, abundant, 280 m, pH 6,5; leg. P. Sunding (XI/1976), 0 (CV-24).


FIG. 5. Ramalinetum maderensis Follm. et Mies with dominating Ramalina rubrotincta Krog et 0sth. and Lecanora sulphurella Hepp on lava face in the middle mountain belt of Sao Nicolau (1986).

40. Ramalina rubrotincta Krog et Osth. - Ramalinaceae This corticolous or saxicolous, acidophytic, endemic thamnolichen of the Cape

Verde Islands has been described originally from Sal (type) and Sao Vicente (Krog & 0sthagen 1978). It is new to Brava, Fogo, Santo Antao, Sao Nicolau, and Sao Tiago ; further collections from Sao Vicente are omitted here. Due to a superficial similarity, Cape Verdean collections of this epilithophyte have formerly been mistaken for Ramatina arabum (Dill .) Mey. et Flot. (Sao Vicente: Bouly de Lesdain 1935), which should


be removed from the check-list of Cape Verdean lichens. According to our field notes, it is a common companion species of various epilithic and epiphytic lichen associations of the mountain belt, sometimes forming dense, monotypic mats on perpendicular, maritime rock faces.

Brava: Volcanic rocks, road above Fajao Agua, scattered, 300 m, NW, pH 6,5 ; leg. W. Lobin (X/1979), FR CV-795, KOELN 22 169. - Volcanic rocks, northern slope of Monte Miranda, abundant, 680 m, N , pH 6,0 ; leg. 0. H. Rustan & C. Brochmann (11/1982), 0 (CV-47). Fogo: Volcanic outcrops, path from Monte Velho to Mosteiros, scattered, 500 m, NE, pH 6,0; leg. G . Follmann (III/1986), KOELN 40 009. Sal : Volcanic rocks, eastern slope of Monte Grande, scarce, 200-300 m, NE, pH 6,0 ; leg. P. Sunding (X/1972), 0 (CV-57). - Volcanic rocks, top of Monte Rocha da Salina, scarce, 300 m, NE, pH 6,0; leg. P. Sunding (X/1972), 0 (CV-58). - Volcanic rocks, summit of Monte Grande, abundant, 400 m, NW, pH 6,0; leg. G. Fol1mann (JU/1 986), KOELN 30010. Santo Antilo: Volcanic rocks, pass height between Proto Novo and Ribeira Grande, scattered, 1200 m, NE, pH 6,0; leg. W. Lobin (Xn/1 979), FR CV-899. Silo Nicolau: Dry shrubs, north-eastern slope of Monte Agua de Jose, scattered, 600 m, NE, pH 6,0; leg. G. Fol1mann (III/1986), KOELN 30 011. Silo Tiago: Dry twigs, north-western slope of Pico da Antonia, scattered, 1000 m, NW, pH 6,5; leg. G . Follmann (IV/ 1974), KOELN 22 180.

DISCUSSION

Nineteen of the 40 lichen species documented here are new to the Cape Verde Islands (e.g., Catapyrenium lachneum (Ach.) Sant.), 18 to single islands of the group (e.g., Heterodermia isidiophora (NyJ.) Awas.), and three to the entire Macaronesian floral region (e.g., Ramalina peruviana Ach.). Moreover, various doubtful records could be confirmed (e.g., Dirina massiliensis Our. et Mont.). Excluding some lichen taxa formerly misunderstood (e.g., Ramalina arabum (Dill.) Mey. et Flot.), the lichen inventory of the Cape Verde Islands now comprises 215 items. (A revised check-list will be published elsewhere.) Only few additions to the macrolichen flora are to be expected in future, particularly within the Parmeliaceae, Roccellaceae, and Usneaceae which are actually under study. On the other side, a thorough elaboration of the numerous corticolous and saxicolous microlichens, especially those of mountain sites of difficult accessibility, will increase the lichen number of the Cape Verde Islands considerably.

Cosmopolitan species clearly predominate among the new findings (16 ; e.g. , Cladoniafurcata (Huds.) Schrad.), followed by Mediterranean (12; e.g., Psora tabacina (Ram.) de Cand.), Tropical (8; e.g., Parmotrema reticulatum (TayJ.) Choisy), and Macaronesian ones (4; e.g., Ramalina bourgaeana Mont.). These observations are not contradictory to the general concept of the chorological tendencies of the Cape Verdean lichen flora (Follmann & S{mchez-Pinto 1986); Tropical geoelements 30%, Mediterranean geoelements 25 %, Cosmopolitan geoelements 20 %, Macaronesian geoelements 15 %, Cape Verdean endemics IO %. Thirty-four of the enumerated lichen taxa occur also on other Macaronesian archipelagos, and 31 of these have been found on the Canary Islands, too. Nevertheless, the lichen colonization of the latter island group with its many Atlantic-Mediterranean (50 %), Boreal-Temperate (15 %), and Arctic-Alpine species (10%) most probably proceeded in a completely different way


TABLE 1. Island distribution of the new lichen records (A = Santo Antao, B = Brava, C = Cape Verde Islands, F = Fogo, I = new record for single islands, M = Macaronesia, N = Sao Nicolau, O = Maio, R = Boa Vista (after Sal-Rei), S= Sal, T = Sao Tiago, V= Sao Vicente, bold face = new record).

Lichen taxa R S N V A 0 T F B

Catapyrenium lachneum (C) ~ Cfadonia fofiacea (l) N A I Cfadonia furcata (C) ~ A I Cfadonia pociflum (C) ~ A Cfadonia rangiformis (l) ~ A T Dipfoschistes scruposus (C) ~ r: Dirina massiliensis (l) § ~ Gorgadesia mira (I ) § ~ V A T B Graphis scripta (C) I Heppia turgida (C) A Helerodermia isidiophora (I) ~ A I F Heterodermia japonica (C) A Leprocaulon microscopicum (l) N Y A T Leptogium furfuraceum (C) V A Leptogium fichenoides (C) A Parmefia borreri (l) A T Parmotrema chinense (l) N V A T Parmotrema reticulatum (l ) ~ V A T r: Peftigera hymenina (M) A Peftigera rufescens (C) A I Pertusaria texana (M) I Phaeophyscia orbicularis (C ) I Physcia tenella (C) ~ Physcia tribacia (C) A Pseudoparmefia caperata (l) N A Psora tabacina (C) ~ Ramafina bourgaeana (l) § N V A Ramalina duriaei (l) S ~ V I B Ramafina macijormis (C) ~ A Ramafina maderensis (l) R § ~ V A Ramalina peruviana (M) § ~ y T Ramafina rubrotincta (l) S ~ V A I r: !! Ramalina subgeniculata (l) ~ V A I Roccella africana (C) R § ~ Roccella allorgei (C) R § N Squamarina cartifaginea (l) A T Teloschistes vil!osus (C) § ~ Tornabea atlantica (l) § ~ A T Usnea baylei (l) ~ y X anthoparmelia tinctina (C) A r:

o. FOLLMANN & B. MIEs: Lichen flora and vegetation of Cape Verde Islands. IV 519

TABLE 2. Main secondary products of selected lichen species from the Cape Verde Islands (I = chemotype I, lI = chemotype IT, H = accumulated in higher concentration, L=accumulated in lower concentration).

Lichen taxa Secondary products Class of compounds

Cladonia foliacea rumarprotocetraric acid (H) depsidone (-)-usnic acid (L) dibenzofuran

Cladonia furcata {atranorin (L) depside fumarprotocetraric acid (H) depsidone

Cladonia pocillum {atranorin (L) depside fumarprotocetraric acid (H) depsidone

ttranorin (H) depside Cladonia rangiformis fumarprotocetraric acid (L) depsidone

(+ )-rangiformic acid (H- aliphatic acid

ttranorin (L) depside Diploschistes scruposus diploschistesic acid (H) depside

lecanoric acid (H) depside

rrythrin (H) depside Dirina massiliensis lecanoric acid (L) depside

portentol (L) aliphatic acid

Gorgadesia mira psoromic acid (H) depsidone

Graphis scripta norstictic acid (H) depsidone

ttranorin (H) depside Heterodermia isidiophora leucotylin (H) triterpenoid

zeorin (L) triterpenoid

ttranorin (H) depside Heterodermia japonica norstictic acid (H) depsidone


ttranorin (H) depside Leprocaulon microscopicum ( - )-usnic acid (L) di benzof uran


Parmelia borreri {atranorin (H) depside gyrophoric acid (H) depside

Parmotrema chinense {atranorin (H) depside stictic acid (H) depsidone

Parmotrema reticulatum {a tranorin (H) depside salazinic acid (H) de psi done

(constictic acid (L) depsidone Pertusaria texana 1 stictic acid (H) depsidone

thiophanic acid (L) xanthone

Phaeophyscia orbicularis skyrin (L) anthraquinone

Physcia tenella atranorin (H) depside

rtranorin (H) depside Physcia tribacia leucotylin (L) triterpenoid

zeorin (H) triterpenoid

rtranorin (L) depside

Pseudoparmelia caperata caperatic acid (H) aIiphatic acid protocetraric acid (L) depsidone ( + )-usnic acid debinzofuran

roUrgeaniC acid (H) aliphatic acid

Ramalina bourgaeana (I) salazinic acid (H) depsidone triterpenoids (L) triterpenoids ( + )-usnic acid (L) dibenzofuran


TABLE 2. (continued)

Lichen taxa Secondary products Class of compounds

rourgeaniC acid (H) aliphatic acid Ramalina bourgaeana (II) triterpenoids (L) triterpenoids

( + )-usnic acid (L) dibenzofuran

Ramalina duriaei {bourgeanic acid (H) aliphatic acid ( + )-usnic acid (L) dibenzofuran

toUrgeaniC acid (H) aliphatic acid Ramalina maci/ormis (I) norstictic acid (H) depsidone

(+ )-usnic acid (L) dibenzofuran

tourgeaniC acid (H) aliphatic acid Ramalina maciformis (11) salazinic acid (H) depsidone


Ramalina maderensis (I) eecanoric acid (H) depside (+ )-usnic acid (L) dibenzofuran

Ramalina maderensis (II) {4-0-demethylbarbatic acid (H) depside ( + )-usnic acid (L) dibenzofuran

(homosekikaic acid (L) depside Ramalina peruviana (sekikaic acid (H) depside


tonnorstictic acid (L) depsidone Ramalina rubrotincta norstictic acid (H) depsidone


rivaricatic acid (H) depside

Ramalina subgeniculata norstictic acid (L) depsidone salazinic acid (L) depsidone (+ )-usnic acid (L) dibenzofuran

rrythrin (H) depside Roccella a/ricana lecanoric acid (L) depside

salazinic acid (L) depsidone

Roccella allorgei {erythrin (H) depside norstictic acid (L) depsidone

Squamarina cartilaginea {psoromic acid (H) depsidone ( - )-usnic acid (L) dibenzofuran

Teloschistes villosus parietin (H) anthraquinone

Usnea baylei {norstictic acid (H) depsidone ( + )-usnic acid (L) dibenzofuran

ttranorin (H) depside Xanthoparmelia tinctina salazinic acid (H) depsidone

( + )-usnic acid (L) dibenzofuran

(Hern{mdez-Padron et al. 1986). Naturally, the larger, higher, and moister islands like Santo Antao show a higher

species diversity than do smaller, lower, and drier ones like Maio. The seemingly greater species density of Santo Antao, Sao Tiago, and Sao Vicente can be explained by easier access and better exploration in former times. Table 1 clearly demonstrates that the relatively humid island of Sao Nicolau, which received more attention recently, bears a similarly rich lichen flora. As a rule, the xerophytic coastal belt of the Cape Verdes is characterized by a fairly uniform lichen population, whilst the hygrophytic mountain belt may present considerable differences from island to island.

G. FOLLMANN & B. MIES : Lichen flora and vegetation of Cape Verde Islands. IV 521

Various lichen-rich plant associations of the Cape Verdes are habitually, floristically, and ecologically similar to those found on other Macaronesian archipelagos, especially the Canaries. If they possess a higher percentage of joint species but are clearly separated by single differential species, they are provisionally interpreted here as local subassociations. The basis associations may have Macaronesian, Mediterranean, or Pantropical origin (e.g., Roccelletum canariensis Follm., Cladonietum foliaceae Klem., Teloschistetum flavicantis Follm., respectively). Other lichen associations show a high degree of independence (e.g., Ramalinetum maderensis Follm. et Mies). A detailed survey of the lichen vegetation of the Cape Verde Islands will be published elsewhere.

No new distribution patterns of secondary lichen products could be observed, but most samples of the Cape Verdean lichens analyzed accumulate comparatively high concentrations of specific compounds (tab. 2). Again, this corresponds to the situation on the Canary Islands (Hernandez-Padron et al. 1986). In view of the many biological and ecological effects demonstrated or postulated for lichen substances (Follmann 1969, Rundel 1978), it can be admitted that higher levels of secondary metabolites increase the competitiveness and viability of pioneer lichens, particularly on younger lava deposits. However, this certainly holds good only for warmer climates because many species of wider distribution show a notable decrease in the formation of such compounds towards the polar regions.

ACKNOWLEDGMENTS

The authors are much indepted to the directors and curators of the herbaria at Francfort (FR), Kassel (KASSEL), Oslo (0), and Santa Cruz de Tenerife (TFMC) for the loan of lichen samples in their care, especially to Dr. H. Krog (Oslo), Dr. W. Lobin (Francfort), Dipl.BioI. L. Sanchez-Pinto (Santa Cruz de Tenerife), and Dr. P. Sunding (Oslo). Moreover, the senior author gratefully acknowledges receipt of a travel grant of the University of Cologne (1986).

SUMMARY

Forty lichen taxa are reported from the Cape Verde Islands and discussed concerning their taxonomy, chorology, ecology, and sociology. Peltigera hymenina (Ach.) Del., Pertusaria texana Muell.Arg., and Ramalina peruviana Ach. represent first records for Macaronesia. Nineteen lichen species are new to the Cape Verde Islands and 18 to single islands of the archipelago. Cosmopolitan geoelements clearly predominate among the new findings (16), followed by Mediterranean (12), Tropical (8), and Macaronesian (4) ones. Excluding some taxa formerly misunderstood, the lichen inventory of the Cape Verde Islands now comprises 215 items. Many of the Cape Verdean lichens accumulate specific secondary products in comparatively high concentrations. This might increase their competitiveness and viability, especially on younger lava deposits . Besides endemical ones, various lichendominated pioneer associations of the Cape Verde Islands are floristically related to those of Pantropical, Mediterranean, or Macaronesian distribution .


ZUSAMMENFASSUNG

Vierzig Flechtenarten werden erstmals von den Kapverden gemeldet und hinsichtlich ihrer Taxonomie, Chorologie, L>kologie und Soziologie besprochen. Peltigera hymenina (Ach.) Del., Pertusaria texana Muell.-Arg. und Ramalina peruviana Ach. stellen Neufunde fUr die makaronesische Florenregion dar; 19 sind Erstfunde fUr die Kapverden, 18 fUr deren Einzelinseln. Unter den Erstnachweisen herrschen kosmopolitische Florenelemente i. w. S. vor (16); es folgen mediterrane (12), tropische (8) und makaronesische (4). Nach Ausschlu13 einiger friiher mil3interpretierter Sippen umfa13t der Flechtenkatalog der Kapverden jetzt 215 Lichenentaxa. Zahlreiche kapverdische Flechtenarten speichern spezifische sekundare Inhaltsstoffe in ungewohnlich hoher Konzentration. Moglicherweise besitzen sekundarstoffreichere Chemotypen hohere Konkurrenzkraft und bessere Ausbreitungsmoglichkeiten als sekundarstoffarmere, besonders auf jungem Ergu13gestein. Neben endemischen finden sich verschiedene flechtenreiche Pioniergesellschaften mit engen floristischen Verwandtschaftsbeziehungen zu solchen der Pantropis, Mediterraneis und Makaronesiens.

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