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Notes on African Pannariaceae (lichenized ascomycetes)
Lichenologist 35(1): 11–20 (2003) doi:10.1006/lich.2002.0424
Notes on African Pannariaceae (lichenized ascomycetes) Per M. JØRGENSEN Abstract: The African continent is shown to contain only 38 species in the lichen family Pannariaceae, all of which are listed in the conclusion. Four new species are described: Pannaria planiuscula (Republic of South Africa [RSA] and Kenya), Pannaria squamulosa (RSA), Parmeliella dactylifera (RSA), and Parmeliella triptophylloides (Kenya). Four species are recorded as new to the continent: Pannaria centrifuga P.M. Jørg. (RSA), Pannaria ramosii Vain. (Tanzania), Parmeliella imbricatula (Müll. Arg.) P. M. Jørg. (RSA), and Psoroma fruticulosum James & Henssen (RSA). The following taxa described from Africa prove to be synonyms: Pannaria cameroonensis Dodge (=Parmeliella stylophora), Pannaria capensis Stirt. (=P. lurida), Pannaria leucosticta var. isidiopsis Nyl. (=P. globigera), Pannaria pityrella Stirt. (=Coccocarpia stellata), and Pannaria thoroldii Dodge (=Parmeliella mariana). Three species have been incorrectly recorded from Africa: Pannaria fulvescens, Parmeliella nigrocincta and Parmeliella triptophylla. 2003 The British Lichen Society. Published by Elsevier Science Ltd. All rights reserved.
Key words: Africa, lichens, new records, new species, taxonomy.
Introduction In my revisions of lichens in the family Pannariaceae, Africa has hitherto been under-represented. I have never visited the African continent, and must rely on material collected by others, which, however, has given me some insight into its pannariaceous flora, reported below. The first survey of African lichens, including the Pannariaceae, is that of Stizenberger (1890). It contains 15 species, a number which surprisingly has only been doubled since that time, mainly with records from East Africa (Swinscow & Krog 1988). Collections from West Africa (except Macaronesia, see Hafellner 1995) are curiously rare, and most possibly underrepresented. Taxa additional to these major surveys are recorded here.
New species and records 1. Moelleropsis nebulosa (Hoffm.) Gyeln. Rabenhorst Krypt. Fl. 9,2: 257 (1940). New Record: Republic of South Africa: Transvaal: Magoeba’s Kloof, terricolous on road-cutting, in open shade, locally abundant, 1946, E.A.C.L. Schelpe 1630 (BG, LD, BOL!).
This is a species readily recognized by its bluish, granular thallus (Jørgensen 2001a). It was previously known only from North Africa (Stizenberger 1890: 85) where it is a Mediterranean element, and an addition to that disjunct element in South Africa (see discussion below). 2. Pannaria centrifuga P. M. Jørg. Bibl. Lich. 78: 113 (2001).
P. M. Jørgensen: Department of Botany, University of Bergen, Allégaten 41, N-5007 Bergen, Norway. 0024–2829/03/010011+10 $30.00/0
New record: Republic of South Africa: Cape Prov.: Western Cape, Kirstenbosch Botanical Garden, on rocks on SE-facing slope with evergreen, subtropical forest, 1996, L. Tibell 21193 (UPS!).
2003 The British Lichen Society. Published by Elsevier Science Ltd. All rights reserved.
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F. 1. Pannaria planiuscula, marginal parts of holotype (L).
This species, recently described from Australia, forms flat, radiating saxicolous rosettes. In the South African material these are rather small (young), up to 3 cm diam., lacking the characteristic central lobules which develop with age, but with some of these along the lobe margins. Otherwise the specimens are in accord with Australian specimens. It is a rather unexpected addition to the flora element which is shared between Australia and South Africa (Rogers & Stevens 1981), since most of these occur in drier habitats (Jørgensen 2002b). Exceptions, however, are known, for example Collema leucocarpum Hook.fil. & Taylor (Rogers 1992: 39), but most of these are also known in western Australia, where P. centrifuga is not known to occur. It is, however, rather widespread in the East, and also present in New Zealand and Tasmania (Jørgensen 2001b).
3. Pannaria planiuscula P. M. Jørg. sp. nov. Pannariae rubiginosae similis sed thallo planissimo, PD non reagens, et sporis minutis. Typus: Kenya, Rift Valley Prov., Trans Nzoia Distr., Cherangani Mts., on tree near rivulet in densely wooded ravine, shaded, 2500–2700 ft alt., 22 May 1949, R. Maas Geestranus 11484 (L—holotypus).
(Fig. 1) Thallus subcrustose, flattened, in rosettes to 5 cm diam., PD, with radiating marginal lobes, up to 2 mm broad, 150–200 m thick with distinct, paraplectenchymatous upper cortex, 30–40 m wide; prothallus inconspicuous; upper surface greyish blue. Central parts small-squamulose to lobulate with numerous apothecia. Apothecia up to 1 mm diam. with distinct thalline margin and brownish disc. Hymenium c. 100 m high, I+ blue only in the vicinity of asci which have no apical
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F. 2. Pannaria ramosii, detail of African specimen (O).
amyloid structures; ascospores simple, colourless, ellipsoid, 10–125–6 m. Pycnidia not observed.
ate altitudes. As yet known only from two disjunct localities in Kenya and South Africa, suggesting a widespread species in this region.
Chemistry. All reactions negative, no substances detected by TLC.
Additional material studied. Republic of South Africa: Cape Prov.: Cape Distr., slope of Table Mt., Blinkwater Ravine, on Eucalyptus, 1953, O. Almborn 1874 & 1880 (LD).
Notes. Pannaria planiuscula may be mistaken for the Indian P. complanata P. M. Jørg., another flat representative in the P. rubiginosa group. Pannaria planiuscula differs in having less complanate marginal lobes not resting on a thick prothallus, and in being PD and with distinctly smaller spores, those of P. complanata being 14– 207–10 m. In these characters P. planiuscula is very close to the Himalayan P. emodi P. M. Jørg., which is a distinctly different species due to its brownish, squamulose thallus. Habitat and distribution. A corticolous species known from moist ravines at moder-
4. Pannaria ramosii Vain. Ann. Acad. Sci. Fenn. ser. A, 15,6: 16 (1920); type: Philippines, Luzon, Prov. Rizal, Aug. 1911, M. Ramos (TUR-V 12186A—lectotype! fide Jørgensen 2001b). New records: Tanzania: Tanga: West Usambara Mts., elfin woodland E ridge of Kilimandege summit in Balangi West Forest Reserve, H. Krog 88208/3-6 (BM, O!); Amani, in the surroundings of Forestry House, in rain forest, 900 m, 1971, R.Moberg 1467a & R. Santesson 23144 (UPS!). Eastern Prov.: Uluguru Mts., above Morogoro, NE ridge of Bondwa between Morningside and Mwere valley, Podocarpus forest, 1750 m, 1972, T. Pocs 6537 (BM!).
(Fig. 2)
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F. 3. Pannaria squamulosa, detail of holotype (UPS).
This long forgotten and misunderstood species was recently treated by Jørgensen (2001b). It may be confused with Pannaria multifida P. M. Jørg. in ed. from the Mascarenes which has repeatedly divided lobules and does not contain pannarin, contrary to P. ramosii which is Pd+ orange (pannarin) with irregular lobules. It is obviously rare in East Africa and confined to elfin, low montane rainforests in the Usambara and associated mountains (the Eastern Arc), which contain a number of relic species (Iversen 1991). Pannaria ramosii also occurs on the Mascarenes, like several other lichens in this region of East Africa (Krog 2000: 17), though it has a wider paleotropical distribution also occurring in S India, the Philippines and NE Australia (Jørgensen 2001b), thus being a widespread paleotropical species of some antiquity.
5. Pannaria squamulosa P. M. Jørg., sp. nov. Pannariae complanatae similis, sed thallo udique squamuloso (sine lobis marginalis elongatis), sine acidis lichenosis, et sporis minutis. Typus: Republic of South Africa, Cape Prov., Knysna Div., Diepvalle Forest Reserve, more or less solitary trees of Cunonia capensis, 5 February 1970, G. Degelius SA-226 (UPS—holotypus).
(Fig. 3) Thallus effuse, up to 4 cm diam., consisting of incised, grey-blue to olivaceus squamules, up to 3 mm wide, 150 m thick, with paraplectenchymatous upper cortex; resting on a well-developed, crustaceous, blackish prothallus. Apothecia common, particularly in central parts of the thallus, up to 1·5 mm diam with plane, brown disc and squamules, greybrown thalline margin, up to 150 m thick.
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Hymenium c. 100 m high, I+ blue in the vicinity of the asci which are without internal amyloid structures, 8-spored; ascospores colourless, simple, subglobose, (10–) 12– 158–10 m, smooth. Pycnidia not observed. Chemistry. All reactions negative, no lichen acids found by TLC. Notes. Pannaria squamulosa closely resembles the recently recognized paleotropical species P. complanata (Jørgensen 2001c) which differs in having elongated, marginal lobes that are PD+ orange (pannarin). The lack of lichen substances may indicate a relationship with Parmeliella mariana (Fr.) P. M. Jørg. & D. J. Galloway, although that species has asci with an amyloid ringstructure, as well as marginal lobes. Another superficially rather similar species is the likewise squamulose, South American Pannaria coeruleonigricans Müll. Arg. which has a more prominent, fibrous prothallus, and contains pannarin. Pannaria squamulosa is certainly most closely related to P. planiuscula which differs markedly in its radiating marginal lobes. Habitat and distribution. A corticolous species as yet known only from the forests in the Cape Province in South Africa. It remains to be seen whether it belongs in the distinctive Cape element (Almborn 1966) or has a wider afroalpine distribution. Additional material studied. Republic of South Africa: Cape Prov.: Ysternek Nat. Park, lower part, 2 km NE of Diepvalle, alt. 450 m, 1996, A. Nordin 4613 (UPS).
6. Parmeliella dactylifera P. M. Jørg., sp. nov. Parmeliellae stylophorae similis, sed thallo minute squamuloso, isidiis dactyliferis apicalibus nigrescentibus; sine acidae lichenosae; sterilis. Typus: Republic of South Africa, Transvaal, Lydenburg distr., 25 km E of Lydenburg, Long Tom Pass at Whisky Spruit, small grove of planted Eucalyptus, 19 February 1984, O. Almborn & I. Kärnefelt 8481—63 (LD—holotypus).
(Fig. 4)
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Thallus small-squamulose, grey-brown, partly variegated, forming colonies up to 5 cm diam., resting on a thin blackish prothallus; individual squamules to 3 mm diam with mainly marginal, digitate isidia up to 1·5 mm long, with enlarged apices that often turn blackish. Thallus 200–250 m thick with a distinct paraplectenchymatous cortex, 25–30 m thick. Apothecia not known fully mature, with indistinct thalline margin and brown disc. Hymenium I+ persistently blue; asci with amyloid apical structures. Ascospores not fully developed, but simple and colourless. Pycnidia not observed. Chemistry. All reactions negative, no substances detected by TLC. Notes. Parmeliella dactylifera has some similarity to Parmeliella stylophora, but is readily distinguished by its small squamulose thallus and a lack of enlarged marginal lobes, as well as its characteristically digitate isidia. It may be the isidiate counterpart of Parmeliella imbricatula. Habitat and distribution. A corticolous species, as yet known only from two localities in South Africa. In the type locality, a particularly humid (‘oceanic’) habitat, it occurs with several other Pannariaceae that are known to be widespread: Pannaria conoplea, P. rubiginosa, P. tavaresii. Additional material studied. Republic of South Africa: Cape Prov.: Table Mt., the southern plateau, 1984, O. Almborn & I. Kärnefelt 84721, 84745, 84753 (LD). Transvaal: Lydenburg Distr., 25 km E. of Lydenburg, Long Tom Pass at Whisky Spruit, 1984, O. Almborn & I. Kärnefelt 8481-64, 8490-68 (LD).
7. Parmeliella imbricatula (Müll. Arg.) P. M. Jørg. ined. Pannaria imbricatula Müll. Arg., Flora 64: 507; type: Brazil, São Paulo, Apiahy, Puiggari 148 p.p. (G— holotypus!). New record: Republic of South Africa: Cape Province: Tzitzikama Mts., Plaat Forest near Storms River, alt. 200 m, 1949, R. A. Maas Gesteeranus 6633 (L).
For a description see Jørgensen (2003). This is a rather unexpected discovery of a species otherwise known only from a few
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F. 4. Parmeliella dactylifera, detail of holotype (LD), showing the characteristic isidia.
collections in S America, indicating a much wider distribution in subtropical regions. 8. Parmeliella triptophylloides P. M. Jørg. sp. nov. Parmeliellae triptophyllae similis sed thallo contiguo et hypothecio apotheciorum incolorato. Typus: Kenya, Rift Valley Prov., Trans Nzoia Distr., Cherangi Mountains, in densely wooded ravine, shaded, c. 900 m, 22 May 1949, R. A. Maas Geesteranus 11483 (L—holotypus; BG, LD—isotypi).
(Fig. 5) Thallus crustose-squamulose, resting on a blackish, crustose prothallus, irregular, covering considerable areas of the branches on which it grows, cracking into areolate parts, individual squamules to 1·5 mm diam., round to elongate, grey-brown, smooth, shallowly incised, with marginal grey-blue coralloid to phyllidiate isidia that eventually cover and obscure the thallus which is 40–60 m thick with distinct paraplectenchymatous upper cortex, 10–15 m wide.
Apothecia rare, up to 1 mm broad, often deformed or proliferating, brown with paler proper exciple, up to 50 m broad; subhymenium of intricately interwoven hyphae, colourless, to 150 m high centrally. Hymenium I+ persistently deep blue; asci with apical amyloid ring-structure. Ascospores simple, hyaline, no mature examples observed, young spores broadly ellipsoid. Pycnidia unknown. Chemistry. All reactions negative, no substances detected by TLC. Notes. Superficially rather like P. triptophylla, but forming bluish, contiguous, though cracking, colonies. Easily distinguished under the microscope by the colourless subhymenium of the apothecia, compared to the distinctly brown subhymenium in P. triptophylla (see Fig. 46D in Jørgensen 1978: 89).
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F. 5. Parmeliella triptophylloides, detail of fertile part of holotype.
Habitat and distribution. As yet known only from the rich type collection which is corticolous on smooth bark. It appears to be a species of humid, rather low altitude hills; localities which according to Swinscow & Krog (1988: 6) are characterized by their richness in lichens containing cyanobacteria. 9. Psoroma fruticulosum P. James & Henssen
the disjunction between New Zealand (from where it was described) and southern South America (with S Georgia & S Shetland Isles; see Øvstedal & Lewis Smith 2001). Discovery of this Psoroma in the mountains of South Africa is not unexpected, for this genus, with a mainly subantarctic distribution, has some species which reach alpine localities of the adjacent continents.
Mycotaxon 18: 31 (1983). New record: Republic of South Africa: Transvaal: Tulbagh Div., Schneeuwgat Pk., mossy, sandy bank on cliffs, 6000 ft, E. Esterhuysen 9419 (BG, LD, PRE).
For a detailed description see Henssen et al. (1983). This species belongs in the Psoroma hypnorum group, and is characterized by the erect isidioid lobules, which are partly flattened. It is new to Africa and the record fills
Excluded or uncertain species This list mainly contains taxa recorded in the above-mentioned works: Pannaria cameroonensis Dodge. The holotype specimen (from Bipinde in Cameroon, leg. Zenker, FH!) of this species, which has hardly been mentioned after its first publication (Dodge 1964), has proven to be a specimen of the variable Parmeliella stylophora Vain.
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Pannaria capensis J. Steiner. Studies of the holotype (South Africa, Mt. Mamoutsuiri, Junoud, W!) have shown it to belong in P. lurida s. lat., a most variable tropical complex (Jørgensen 2001a), to which it should be referred. A further, better developed specimen (Cape, Forest Hall, A. Duthie 416, BOL) confirms this. Pannaria fulvescens (Mont.) Nyl. s.str. is a species confined to the South Pacific. As shown by Jørgensen & Kashiwadani (2001), the specimens so named in East Africa belong in P. globuligera Hue, which is now also known to occur as far south as the Cape region (Table Mt., NE side, E. Esterhuysen 8912, LD). Pannaria leucolepis Wahlenb. This name is a synonym of Pannaria hookeri, a species known from only one collection from high altitudes in East Africa (Frisch & Hertel 1998). Stizenberger’s record is a MacOwan specimen from Table Mountain in the Cape of Good Hope, which I have not seen, but possibly represents an extreme form of P. rubiginosa, which is recorded from that region. Pannaria leucosticta Tuck. This is now known as Fuscopannaria leucosticta (Tuck.) P. M. Jørg., a widespread species in warmtemperate to subtropical regions of America and East Asia, which once also occurred in Mediterranean Europe (Jørgensen 1978). The specimen from Madeira (Mandon 37) is not that species, but the closely related F. mediterranea (C. Tav.) P. M. Jørg. Fuscopannaria leucosticta is not known with certainty to occur in Africa. var. isidiopsis Nyl. in Cromb. is known only from the small holotype specimen from Table Mountain in the Cape of Good Hope (leg. Eaton, BM!). This is an unusual expression of Pannaria globuligera Hue, a species with a wide distribution in East Africa, rather than P. tavaresii as recorded by Jørgensen & Galloway (1992: 270), a species which is also present in the region (Cape, Table Mt., lower plateau, E. Esterhuysen 20100, LD). Pannaria pityrella Stirt., a species which when originally described from Nigeria (Bonny River), was compared with Pannaria asterella Nyl. (=Coccocarpia stellata Tuck.).
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Studies of the holotype (GLAM), have shown that it is just a form of this quite variable species (Arvidsson 1982: 88). Coccocarpia stellata is a new record for West Africa of this predominantly neotropical species. Pannaria thoroldii Dodge is, like P. cameroonensis, described from a specimen from Cameroon and has not been recollected since its description (Dodge 1964). The holotype specimen (Thorold 2342, FH!) proves to belong in Parmeliella mariana s.lat., a complex in need of further revision, though this specimen does not stand out as something worth separate taxonomic recognition. Parmeliella nigrocincta (Mont.) Müll. Arg. is not known in its strict sense from Africa. Records from the Canary Islands are mainly based on Parmeliella miradorensis Vain. (Jørgensen 1998), and those from South and East Africa are Parmeliella coelestina Zahlbr. Parmeliella triptophylla (Ach.) Müll. Arg. is not known with certainty from the African continent. Records from the Cape may be P. lacerata P. M. Jørg. (Jørgensen 1998) or those from further north P. triptophylloides P. M. Jørg. It is, however, present in Macaronesia (Hafellner 1995). Checklist of African Pannariaceae The following Pannariaceae (38) are now known to occur on the African continent: Degelia atlantica (Degel.) P. M. Jørg. & P. James (Tunisia, Jørgensen 1978: 20). Degelia plumbea (Lightf.) P. M. Jørg. & P. James (North Africa, Jørgensen 1978: 57). Erioderma leylandii (Taylor) Müll. Arg. (Republic of South Africa, Jørgensen 2001d; Tanzania, Krog 2000: 23). Erioderma meiocarpum Nyl. (Ethiopia, East Africa, Rwanda, Zambia, Swinscow & Krog 1994: 84). Erioderma mollissimum (Samp.) Du Rietz (Kenya, Jørgensen & Arvidsson 2001: 508). Erioderma sorediatum D. J. Galloway & P. M. Jørg. (Tanzania, Krog 2000: 23; Jørgensen & Arvidsson 2001: 511). Erioderma unguigerum (Bory) Nyl. (Tanzania, Jørgensen 2001d).
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Erioderma verruculosum (Vain.) Hue (Malawi & Zambia, Jørgensen 2001d). Fuscopannaria ignobilis (Anzi) P. M. Jørg. (NW Africa & Kenya, Jørgensen 1978: 33). Fuscopannaria mediterranea (C. Tav.) P. M. Jørg. (NW Africa, Jørgensen 1978: 46). Fuscopannaria olivacea (P. M. Jørg.) P. M. Jørg. (NW Africa, Jørgensen 1978: 51). Fuscopannaria praetermissa (Nyl.) P. M. Jørg. (Kenya, Frisch 1999: 83). Fuscopannaria sampaiana (C. Tav.) P. M. Jørg. (Tunisia, Jørgensen 1978: 65). Fuscopannaria subimmixta (C. Knight) P. M. Jørg. (Republic of South Africa, Jørgensen 2002a: 328). Kroswia crystallifera P. M. Jørg. (Republic of South Africa & Kenya, Jørgensen 2002b) Moelleropsis nebulosa (Hoffm.) Gyeln. (Morocco, Stizenberger 1890: 85; Republic of South Africa). Pannaria centrifuga P. M. Jørg. (Republic of South Africa). Pannaria conoplea (Ach.) Bory (East Africa, Swinscow & Krog 1988: 152; Republic of South Africa, Stizenberger 1890: 84). Pannaria hookeri (Borr.) Nyl. (Kenya, Frisch & Hertel 1998: 368). Pannaria elatior Stirt. (East Africa, Krog 2000 p.p.). Pannaria globigera Hue (Republic of South Africa to Ethiopia). Pannaria lurida (Mont.) Nyl. (Republic of South Africa & East Africa, Swinscow & Krog 1988: 153). Pannaria planiuscula P. M. Jørg. (Kenya & Republic of South Africa). Pannaria rubiginosa (Ach.) Bory (Republic of South Africa to Ethiopia, Swinscow & Krog 1988: 154). Pannaria santessonii Swinscow & Krog (Tanzania, Swinscow & Krog 1988: 155). Pannaria squamulosa P. M. Jørg. (Republic of South Africa). Pannaria tavaresii P. M. Jørg. (Republic of South Africa to Ethiopia, Krog 2000). Parmeliella coelestina Zahlbr. (Republic of South Africa, Jørgensen 1998). Parmeliella dactylifera P. M. Jørg. (Republic of South Africa). Parmeliella furfuracea P. M. Jørg. (Republic of South Africa, Jørgensen 2001c).
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Parmeliella imbricatula (Müll. Arg.) P. M. Jørg. (Republic of South Africa). Parmeliella lacerata P. M. Jørg. (Republic of South Africa, Jørgensen 1998). Parmeliella mariana (Fr.) P. M. Jørg. & D. J. Galloway (East Africa, Swinscow & Krog 1988: 154; Cameroon). Parmeliella stylophora Vain. (Tanzania, Swinscow & Krog 1988: 156; Cameroon). Parmeliella triptophylloides P. M. Jørg. (Kenya). Protopannaria pezizoides (Weber) S. Ekman & P. M. Jørg. (Algeria & Uganda, Jørgensen 1978: 53). Psoroma asperellum Nyl. (Republic of South Africa, Nylander 1863: 24). Psoroma fruticulosum P. James & Henssen (Republic of South Africa). There are additional taxa on some of the islands [Macaronesia (Hafellner 1995), Madagascar, Seychelles, The Mascarenes & Prince Edward Islands (Jørgensen 2000), St Helena, Ascension & Tristan da Cunha (Jørgensen 1977)], which are not included in the list. Discussion The Pannariaceae flora of the mountains in NW Africa (The Atlas and connected chains) is closely linked to that of Macaronesia and SW Europe, but is poorer in taxa. A Mediterranean element can also be traced in other parts of the continent. Well-known is the connection with the distant Cape Province (Almborn 1966), to which the presence of Moelleropsis nebulosa is an addition. More remarkable is the presence of Fuscopannaria ignobilis in East Africa, a region not particularly well-known for its Mediterranean elements, though flowering plants are known to have penetrated south through Egypt and Sudan, and this appears to be a possible route for the taxa reaching as far south as the Republic of South Africa. The East African mountains also contain an arctic-alpine element, for example Fuscopannaria praetermissa, Pannaria hookeri and Protopannaria pezizoides, all at high altitudes (d4000 m). Not unexpectedly, East Africa with its high mountains and varied habitats (Swinscow & Krog 1988) has the richest
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Pannariaceae biota, ranging from the Republic of South Africa to Ethiopia, mostly containing pan(sub)tropical species, a few with Indian connections (Erioderma meiocarpum, Kroswia crystallifera), and some species otherwise restricted to South America (e.g. Erioderma verruculosum) or Australasia (Fuscopannaria subimmixta, Pannaria centrifuga and Parmeliella furfuracea)—the last three in the Republic of South Africa (for further discussion of this element, see Jørgensen 2002b). There is also a small subantarctic element in the mountains of South Africa (Psoroma), which have a varied pannariaceous flora with additional afroalpine, Mediterranean, and subtropical elements, as well as possible endemic taxa. The scarcity of the Pannariaceae in West Africa (excluding Macaronesia) is remarkable, and possibly caused by the scarce collecting in these parts of the continent. In general, Africa has remarkably few species in this lichen family compared with Australasia and South America, which appear to contain the evolutionary centres for several of the genera. The African species, on the other hand, give, with few exceptions, the impression of being remnants of once more widespread taxa. R Almborn, O. (1966) Revision of some lichen genera in South Africa. Botaniska Notiser 119: 73–112. Arvidsson, L. (1982) A monograph of the lichen genus Coccocarpia. Opera Botanica 67: 1–96. Dodge, C. W. (1964) Some lichens of Tropical Africa IV. Beiheft Nova Hedwigia 12: 1–225. Frisch, A. (1999) Afroalpine macrolichens of Mount Kenya. Bayreuther Forum Ökologie 64: 65–102. Frisch, A. & Hertel, H. (1998) Flora of macrolichens in the alpine and subalpine zones of Mount Kenya (Kenya). Sauteria 9: 363–370. Hafellner, J. (1995) A new checklist of lichens and lichenicolous fungi of insular Laurimacaronesia, including a lichenological bibliography of the area. Fritschiana 5: 1–132. Henssen, A. M., Renner, B., Marton, K., James, P. W. & Galloway, D. J. (1983) Studies in the lichen genus Psoroma 2. Psoroma fruticolosum and P. rubromarginatum. Mycotaxon 18: 29–48. Iversen, S. T. (1991) The Usambara mountains, NE Tanzania. Phytogeography of vascular plant flora. Symbolae Botanicae Upsalienses 29(3): 1–234.
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Jørgensen, P. M. (1977) Foliose and fruiticose lichens from Tristan da Cunha. Skrifter fra Det norske Videnskapsakademi i Oslo, Matematisk naturvitenskapelig klasse, n.ser. 36: 1–40. Jørgensen, P. M. (1978) The lichen family Pannariaceae in Europe. Opera Botanica 45: 1–123. Jørgensen, P. M. (1998) Studies in the lichen family Pannariaceae VII. On some poorly known Parmeliella-like species from the Southern Hemisphere. Lichenologist 30: 533–541. Jørgensen, P. M. (2000) Studies in the lichen family Pannariaceae IX. A revision of Pannaria subg. Chryopannaria. Nova Hedwigia 71: 405–414. Jørgensen, P. M. (2001a) Survey of the lichen family Pannariaceae on the American continent, North of Mexico. Bryologist 103: 670–704. Jørgensen, P. M. (2001b) New species and records of the lichen family Pannariaceae from Australia. Bibliotheca Lichenologica 78: 109–139. Jørgensen, P. M. (2001c) Four new Asian species in the lichen genus Pannaria. Lichenologist 33: 297–302. Jørgensen, P. M. (2001d) On the status of the Erioderma names in Zahlbruckner’s Catalogus. Taxon 55: 525–541. Jørgensen, P. M. (2002a) Two species of the Pannariaceae (lichenized ascomycetes) from New Zealand with a wider Southern Hemisphere distribution. New Zealand Journal of Botany 40: 327–329. Jørgensen, P. M. (2002b) Kroswia, a new genus in the Pannariaceae (lichenized ascomycetes). Lichenologist 34: 297–303. Jørgensen, P. M. & Arvidsson, L. (2001) The sorediate species of the lichen genus Erioderma Fée. Nova Hedwigia 73: 497–512. Jørgensen, P. M. & Galloway, D. J. (1992) Pannariaceae. Flora of Australia 54: 246–293. Jørgensen, P. M. & Kashiwadani. (2001) New or misunderstood species of Japanese Pannaria (Lichenes). Journal of Japanese Botany 76: 1–10. Krog, H. (2000) Corticolous macrolichens of low montane rainforests and moist woodlands of eastern Tanzania. Sommerfeltia 28: 1–75. Nylander, W. (1863) Synopsis Lichenum 2. Paris. Øvstedal, D. O. & Lewis Smith, R. (2001) Lichens of Antarctica and South Georgia – A Guide to Their Identification and Ecology. Cambridge: Cambridge University Press. Rogers, R. W. (1992) Lichen ecology and biogeography. Flora of Australia 54: 30–42. Rogers, R. W. & Stevens, N. (1981) Lichens. In Ecological Biogeography of Australia (P. L. Keast, ed.). The Hague: Junk. Stizenberger, E. (1890) Lichenaea africana. Fasc.1. Berichte der Tätigkeit der St. Gallen Naturwissenschaftlichen Gesellschaft 1889/1890: 1–144. Swinscow, T. D. V. & Krog, H. (1988) Macrolichens of East Africa. London: British Museum (Natural History).
Accepted for publication 4 October 2002
Notes on African Pannariaceae (lichenized ascomycetes) Per M. JØRGENSEN Abstract: The African continent is shown to contain only 38 species in the lichen family Pannariaceae, all of which are listed in the conclusion. Four new species are described: Pannaria planiuscula (Republic of South Africa [RSA] and Kenya), Pannaria squamulosa (RSA), Parmeliella dactylifera (RSA), and Parmeliella triptophylloides (Kenya). Four species are recorded as new to the continent: Pannaria centrifuga P.M. Jørg. (RSA), Pannaria ramosii Vain. (Tanzania), Parmeliella imbricatula (Müll. Arg.) P. M. Jørg. (RSA), and Psoroma fruticulosum James & Henssen (RSA). The following taxa described from Africa prove to be synonyms: Pannaria cameroonensis Dodge (=Parmeliella stylophora), Pannaria capensis Stirt. (=P. lurida), Pannaria leucosticta var. isidiopsis Nyl. (=P. globigera), Pannaria pityrella Stirt. (=Coccocarpia stellata), and Pannaria thoroldii Dodge (=Parmeliella mariana). Three species have been incorrectly recorded from Africa: Pannaria fulvescens, Parmeliella nigrocincta and Parmeliella triptophylla. 2003 The British Lichen Society. Published by Elsevier Science Ltd. All rights reserved.
Key words: Africa, lichens, new records, new species, taxonomy.
Introduction In my revisions of lichens in the family Pannariaceae, Africa has hitherto been under-represented. I have never visited the African continent, and must rely on material collected by others, which, however, has given me some insight into its pannariaceous flora, reported below. The first survey of African lichens, including the Pannariaceae, is that of Stizenberger (1890). It contains 15 species, a number which surprisingly has only been doubled since that time, mainly with records from East Africa (Swinscow & Krog 1988). Collections from West Africa (except Macaronesia, see Hafellner 1995) are curiously rare, and most possibly underrepresented. Taxa additional to these major surveys are recorded here.
New species and records 1. Moelleropsis nebulosa (Hoffm.) Gyeln. Rabenhorst Krypt. Fl. 9,2: 257 (1940). New Record: Republic of South Africa: Transvaal: Magoeba’s Kloof, terricolous on road-cutting, in open shade, locally abundant, 1946, E.A.C.L. Schelpe 1630 (BG, LD, BOL!).
This is a species readily recognized by its bluish, granular thallus (Jørgensen 2001a). It was previously known only from North Africa (Stizenberger 1890: 85) where it is a Mediterranean element, and an addition to that disjunct element in South Africa (see discussion below). 2. Pannaria centrifuga P. M. Jørg. Bibl. Lich. 78: 113 (2001).
P. M. Jørgensen: Department of Botany, University of Bergen, Allégaten 41, N-5007 Bergen, Norway. 0024–2829/03/010011+10 $30.00/0
New record: Republic of South Africa: Cape Prov.: Western Cape, Kirstenbosch Botanical Garden, on rocks on SE-facing slope with evergreen, subtropical forest, 1996, L. Tibell 21193 (UPS!).
2003 The British Lichen Society. Published by Elsevier Science Ltd. All rights reserved.
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F. 1. Pannaria planiuscula, marginal parts of holotype (L).
This species, recently described from Australia, forms flat, radiating saxicolous rosettes. In the South African material these are rather small (young), up to 3 cm diam., lacking the characteristic central lobules which develop with age, but with some of these along the lobe margins. Otherwise the specimens are in accord with Australian specimens. It is a rather unexpected addition to the flora element which is shared between Australia and South Africa (Rogers & Stevens 1981), since most of these occur in drier habitats (Jørgensen 2002b). Exceptions, however, are known, for example Collema leucocarpum Hook.fil. & Taylor (Rogers 1992: 39), but most of these are also known in western Australia, where P. centrifuga is not known to occur. It is, however, rather widespread in the East, and also present in New Zealand and Tasmania (Jørgensen 2001b).
3. Pannaria planiuscula P. M. Jørg. sp. nov. Pannariae rubiginosae similis sed thallo planissimo, PD non reagens, et sporis minutis. Typus: Kenya, Rift Valley Prov., Trans Nzoia Distr., Cherangani Mts., on tree near rivulet in densely wooded ravine, shaded, 2500–2700 ft alt., 22 May 1949, R. Maas Geestranus 11484 (L—holotypus).
(Fig. 1) Thallus subcrustose, flattened, in rosettes to 5 cm diam., PD, with radiating marginal lobes, up to 2 mm broad, 150–200 m thick with distinct, paraplectenchymatous upper cortex, 30–40 m wide; prothallus inconspicuous; upper surface greyish blue. Central parts small-squamulose to lobulate with numerous apothecia. Apothecia up to 1 mm diam. with distinct thalline margin and brownish disc. Hymenium c. 100 m high, I+ blue only in the vicinity of asci which have no apical
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F. 2. Pannaria ramosii, detail of African specimen (O).
amyloid structures; ascospores simple, colourless, ellipsoid, 10–125–6 m. Pycnidia not observed.
ate altitudes. As yet known only from two disjunct localities in Kenya and South Africa, suggesting a widespread species in this region.
Chemistry. All reactions negative, no substances detected by TLC.
Additional material studied. Republic of South Africa: Cape Prov.: Cape Distr., slope of Table Mt., Blinkwater Ravine, on Eucalyptus, 1953, O. Almborn 1874 & 1880 (LD).
Notes. Pannaria planiuscula may be mistaken for the Indian P. complanata P. M. Jørg., another flat representative in the P. rubiginosa group. Pannaria planiuscula differs in having less complanate marginal lobes not resting on a thick prothallus, and in being PD and with distinctly smaller spores, those of P. complanata being 14– 207–10 m. In these characters P. planiuscula is very close to the Himalayan P. emodi P. M. Jørg., which is a distinctly different species due to its brownish, squamulose thallus. Habitat and distribution. A corticolous species known from moist ravines at moder-
4. Pannaria ramosii Vain. Ann. Acad. Sci. Fenn. ser. A, 15,6: 16 (1920); type: Philippines, Luzon, Prov. Rizal, Aug. 1911, M. Ramos (TUR-V 12186A—lectotype! fide Jørgensen 2001b). New records: Tanzania: Tanga: West Usambara Mts., elfin woodland E ridge of Kilimandege summit in Balangi West Forest Reserve, H. Krog 88208/3-6 (BM, O!); Amani, in the surroundings of Forestry House, in rain forest, 900 m, 1971, R.Moberg 1467a & R. Santesson 23144 (UPS!). Eastern Prov.: Uluguru Mts., above Morogoro, NE ridge of Bondwa between Morningside and Mwere valley, Podocarpus forest, 1750 m, 1972, T. Pocs 6537 (BM!).
(Fig. 2)
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F. 3. Pannaria squamulosa, detail of holotype (UPS).
This long forgotten and misunderstood species was recently treated by Jørgensen (2001b). It may be confused with Pannaria multifida P. M. Jørg. in ed. from the Mascarenes which has repeatedly divided lobules and does not contain pannarin, contrary to P. ramosii which is Pd+ orange (pannarin) with irregular lobules. It is obviously rare in East Africa and confined to elfin, low montane rainforests in the Usambara and associated mountains (the Eastern Arc), which contain a number of relic species (Iversen 1991). Pannaria ramosii also occurs on the Mascarenes, like several other lichens in this region of East Africa (Krog 2000: 17), though it has a wider paleotropical distribution also occurring in S India, the Philippines and NE Australia (Jørgensen 2001b), thus being a widespread paleotropical species of some antiquity.
5. Pannaria squamulosa P. M. Jørg., sp. nov. Pannariae complanatae similis, sed thallo udique squamuloso (sine lobis marginalis elongatis), sine acidis lichenosis, et sporis minutis. Typus: Republic of South Africa, Cape Prov., Knysna Div., Diepvalle Forest Reserve, more or less solitary trees of Cunonia capensis, 5 February 1970, G. Degelius SA-226 (UPS—holotypus).
(Fig. 3) Thallus effuse, up to 4 cm diam., consisting of incised, grey-blue to olivaceus squamules, up to 3 mm wide, 150 m thick, with paraplectenchymatous upper cortex; resting on a well-developed, crustaceous, blackish prothallus. Apothecia common, particularly in central parts of the thallus, up to 1·5 mm diam with plane, brown disc and squamules, greybrown thalline margin, up to 150 m thick.
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Hymenium c. 100 m high, I+ blue in the vicinity of the asci which are without internal amyloid structures, 8-spored; ascospores colourless, simple, subglobose, (10–) 12– 158–10 m, smooth. Pycnidia not observed. Chemistry. All reactions negative, no lichen acids found by TLC. Notes. Pannaria squamulosa closely resembles the recently recognized paleotropical species P. complanata (Jørgensen 2001c) which differs in having elongated, marginal lobes that are PD+ orange (pannarin). The lack of lichen substances may indicate a relationship with Parmeliella mariana (Fr.) P. M. Jørg. & D. J. Galloway, although that species has asci with an amyloid ringstructure, as well as marginal lobes. Another superficially rather similar species is the likewise squamulose, South American Pannaria coeruleonigricans Müll. Arg. which has a more prominent, fibrous prothallus, and contains pannarin. Pannaria squamulosa is certainly most closely related to P. planiuscula which differs markedly in its radiating marginal lobes. Habitat and distribution. A corticolous species as yet known only from the forests in the Cape Province in South Africa. It remains to be seen whether it belongs in the distinctive Cape element (Almborn 1966) or has a wider afroalpine distribution. Additional material studied. Republic of South Africa: Cape Prov.: Ysternek Nat. Park, lower part, 2 km NE of Diepvalle, alt. 450 m, 1996, A. Nordin 4613 (UPS).
6. Parmeliella dactylifera P. M. Jørg., sp. nov. Parmeliellae stylophorae similis, sed thallo minute squamuloso, isidiis dactyliferis apicalibus nigrescentibus; sine acidae lichenosae; sterilis. Typus: Republic of South Africa, Transvaal, Lydenburg distr., 25 km E of Lydenburg, Long Tom Pass at Whisky Spruit, small grove of planted Eucalyptus, 19 February 1984, O. Almborn & I. Kärnefelt 8481—63 (LD—holotypus).
(Fig. 4)
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Thallus small-squamulose, grey-brown, partly variegated, forming colonies up to 5 cm diam., resting on a thin blackish prothallus; individual squamules to 3 mm diam with mainly marginal, digitate isidia up to 1·5 mm long, with enlarged apices that often turn blackish. Thallus 200–250 m thick with a distinct paraplectenchymatous cortex, 25–30 m thick. Apothecia not known fully mature, with indistinct thalline margin and brown disc. Hymenium I+ persistently blue; asci with amyloid apical structures. Ascospores not fully developed, but simple and colourless. Pycnidia not observed. Chemistry. All reactions negative, no substances detected by TLC. Notes. Parmeliella dactylifera has some similarity to Parmeliella stylophora, but is readily distinguished by its small squamulose thallus and a lack of enlarged marginal lobes, as well as its characteristically digitate isidia. It may be the isidiate counterpart of Parmeliella imbricatula. Habitat and distribution. A corticolous species, as yet known only from two localities in South Africa. In the type locality, a particularly humid (‘oceanic’) habitat, it occurs with several other Pannariaceae that are known to be widespread: Pannaria conoplea, P. rubiginosa, P. tavaresii. Additional material studied. Republic of South Africa: Cape Prov.: Table Mt., the southern plateau, 1984, O. Almborn & I. Kärnefelt 84721, 84745, 84753 (LD). Transvaal: Lydenburg Distr., 25 km E. of Lydenburg, Long Tom Pass at Whisky Spruit, 1984, O. Almborn & I. Kärnefelt 8481-64, 8490-68 (LD).
7. Parmeliella imbricatula (Müll. Arg.) P. M. Jørg. ined. Pannaria imbricatula Müll. Arg., Flora 64: 507; type: Brazil, São Paulo, Apiahy, Puiggari 148 p.p. (G— holotypus!). New record: Republic of South Africa: Cape Province: Tzitzikama Mts., Plaat Forest near Storms River, alt. 200 m, 1949, R. A. Maas Gesteeranus 6633 (L).
For a description see Jørgensen (2003). This is a rather unexpected discovery of a species otherwise known only from a few
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F. 4. Parmeliella dactylifera, detail of holotype (LD), showing the characteristic isidia.
collections in S America, indicating a much wider distribution in subtropical regions. 8. Parmeliella triptophylloides P. M. Jørg. sp. nov. Parmeliellae triptophyllae similis sed thallo contiguo et hypothecio apotheciorum incolorato. Typus: Kenya, Rift Valley Prov., Trans Nzoia Distr., Cherangi Mountains, in densely wooded ravine, shaded, c. 900 m, 22 May 1949, R. A. Maas Geesteranus 11483 (L—holotypus; BG, LD—isotypi).
(Fig. 5) Thallus crustose-squamulose, resting on a blackish, crustose prothallus, irregular, covering considerable areas of the branches on which it grows, cracking into areolate parts, individual squamules to 1·5 mm diam., round to elongate, grey-brown, smooth, shallowly incised, with marginal grey-blue coralloid to phyllidiate isidia that eventually cover and obscure the thallus which is 40–60 m thick with distinct paraplectenchymatous upper cortex, 10–15 m wide.
Apothecia rare, up to 1 mm broad, often deformed or proliferating, brown with paler proper exciple, up to 50 m broad; subhymenium of intricately interwoven hyphae, colourless, to 150 m high centrally. Hymenium I+ persistently deep blue; asci with apical amyloid ring-structure. Ascospores simple, hyaline, no mature examples observed, young spores broadly ellipsoid. Pycnidia unknown. Chemistry. All reactions negative, no substances detected by TLC. Notes. Superficially rather like P. triptophylla, but forming bluish, contiguous, though cracking, colonies. Easily distinguished under the microscope by the colourless subhymenium of the apothecia, compared to the distinctly brown subhymenium in P. triptophylla (see Fig. 46D in Jørgensen 1978: 89).
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F. 5. Parmeliella triptophylloides, detail of fertile part of holotype.
Habitat and distribution. As yet known only from the rich type collection which is corticolous on smooth bark. It appears to be a species of humid, rather low altitude hills; localities which according to Swinscow & Krog (1988: 6) are characterized by their richness in lichens containing cyanobacteria. 9. Psoroma fruticulosum P. James & Henssen
the disjunction between New Zealand (from where it was described) and southern South America (with S Georgia & S Shetland Isles; see Øvstedal & Lewis Smith 2001). Discovery of this Psoroma in the mountains of South Africa is not unexpected, for this genus, with a mainly subantarctic distribution, has some species which reach alpine localities of the adjacent continents.
Mycotaxon 18: 31 (1983). New record: Republic of South Africa: Transvaal: Tulbagh Div., Schneeuwgat Pk., mossy, sandy bank on cliffs, 6000 ft, E. Esterhuysen 9419 (BG, LD, PRE).
For a detailed description see Henssen et al. (1983). This species belongs in the Psoroma hypnorum group, and is characterized by the erect isidioid lobules, which are partly flattened. It is new to Africa and the record fills
Excluded or uncertain species This list mainly contains taxa recorded in the above-mentioned works: Pannaria cameroonensis Dodge. The holotype specimen (from Bipinde in Cameroon, leg. Zenker, FH!) of this species, which has hardly been mentioned after its first publication (Dodge 1964), has proven to be a specimen of the variable Parmeliella stylophora Vain.
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Pannaria capensis J. Steiner. Studies of the holotype (South Africa, Mt. Mamoutsuiri, Junoud, W!) have shown it to belong in P. lurida s. lat., a most variable tropical complex (Jørgensen 2001a), to which it should be referred. A further, better developed specimen (Cape, Forest Hall, A. Duthie 416, BOL) confirms this. Pannaria fulvescens (Mont.) Nyl. s.str. is a species confined to the South Pacific. As shown by Jørgensen & Kashiwadani (2001), the specimens so named in East Africa belong in P. globuligera Hue, which is now also known to occur as far south as the Cape region (Table Mt., NE side, E. Esterhuysen 8912, LD). Pannaria leucolepis Wahlenb. This name is a synonym of Pannaria hookeri, a species known from only one collection from high altitudes in East Africa (Frisch & Hertel 1998). Stizenberger’s record is a MacOwan specimen from Table Mountain in the Cape of Good Hope, which I have not seen, but possibly represents an extreme form of P. rubiginosa, which is recorded from that region. Pannaria leucosticta Tuck. This is now known as Fuscopannaria leucosticta (Tuck.) P. M. Jørg., a widespread species in warmtemperate to subtropical regions of America and East Asia, which once also occurred in Mediterranean Europe (Jørgensen 1978). The specimen from Madeira (Mandon 37) is not that species, but the closely related F. mediterranea (C. Tav.) P. M. Jørg. Fuscopannaria leucosticta is not known with certainty to occur in Africa. var. isidiopsis Nyl. in Cromb. is known only from the small holotype specimen from Table Mountain in the Cape of Good Hope (leg. Eaton, BM!). This is an unusual expression of Pannaria globuligera Hue, a species with a wide distribution in East Africa, rather than P. tavaresii as recorded by Jørgensen & Galloway (1992: 270), a species which is also present in the region (Cape, Table Mt., lower plateau, E. Esterhuysen 20100, LD). Pannaria pityrella Stirt., a species which when originally described from Nigeria (Bonny River), was compared with Pannaria asterella Nyl. (=Coccocarpia stellata Tuck.).
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Studies of the holotype (GLAM), have shown that it is just a form of this quite variable species (Arvidsson 1982: 88). Coccocarpia stellata is a new record for West Africa of this predominantly neotropical species. Pannaria thoroldii Dodge is, like P. cameroonensis, described from a specimen from Cameroon and has not been recollected since its description (Dodge 1964). The holotype specimen (Thorold 2342, FH!) proves to belong in Parmeliella mariana s.lat., a complex in need of further revision, though this specimen does not stand out as something worth separate taxonomic recognition. Parmeliella nigrocincta (Mont.) Müll. Arg. is not known in its strict sense from Africa. Records from the Canary Islands are mainly based on Parmeliella miradorensis Vain. (Jørgensen 1998), and those from South and East Africa are Parmeliella coelestina Zahlbr. Parmeliella triptophylla (Ach.) Müll. Arg. is not known with certainty from the African continent. Records from the Cape may be P. lacerata P. M. Jørg. (Jørgensen 1998) or those from further north P. triptophylloides P. M. Jørg. It is, however, present in Macaronesia (Hafellner 1995). Checklist of African Pannariaceae The following Pannariaceae (38) are now known to occur on the African continent: Degelia atlantica (Degel.) P. M. Jørg. & P. James (Tunisia, Jørgensen 1978: 20). Degelia plumbea (Lightf.) P. M. Jørg. & P. James (North Africa, Jørgensen 1978: 57). Erioderma leylandii (Taylor) Müll. Arg. (Republic of South Africa, Jørgensen 2001d; Tanzania, Krog 2000: 23). Erioderma meiocarpum Nyl. (Ethiopia, East Africa, Rwanda, Zambia, Swinscow & Krog 1994: 84). Erioderma mollissimum (Samp.) Du Rietz (Kenya, Jørgensen & Arvidsson 2001: 508). Erioderma sorediatum D. J. Galloway & P. M. Jørg. (Tanzania, Krog 2000: 23; Jørgensen & Arvidsson 2001: 511). Erioderma unguigerum (Bory) Nyl. (Tanzania, Jørgensen 2001d).
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Erioderma verruculosum (Vain.) Hue (Malawi & Zambia, Jørgensen 2001d). Fuscopannaria ignobilis (Anzi) P. M. Jørg. (NW Africa & Kenya, Jørgensen 1978: 33). Fuscopannaria mediterranea (C. Tav.) P. M. Jørg. (NW Africa, Jørgensen 1978: 46). Fuscopannaria olivacea (P. M. Jørg.) P. M. Jørg. (NW Africa, Jørgensen 1978: 51). Fuscopannaria praetermissa (Nyl.) P. M. Jørg. (Kenya, Frisch 1999: 83). Fuscopannaria sampaiana (C. Tav.) P. M. Jørg. (Tunisia, Jørgensen 1978: 65). Fuscopannaria subimmixta (C. Knight) P. M. Jørg. (Republic of South Africa, Jørgensen 2002a: 328). Kroswia crystallifera P. M. Jørg. (Republic of South Africa & Kenya, Jørgensen 2002b) Moelleropsis nebulosa (Hoffm.) Gyeln. (Morocco, Stizenberger 1890: 85; Republic of South Africa). Pannaria centrifuga P. M. Jørg. (Republic of South Africa). Pannaria conoplea (Ach.) Bory (East Africa, Swinscow & Krog 1988: 152; Republic of South Africa, Stizenberger 1890: 84). Pannaria hookeri (Borr.) Nyl. (Kenya, Frisch & Hertel 1998: 368). Pannaria elatior Stirt. (East Africa, Krog 2000 p.p.). Pannaria globigera Hue (Republic of South Africa to Ethiopia). Pannaria lurida (Mont.) Nyl. (Republic of South Africa & East Africa, Swinscow & Krog 1988: 153). Pannaria planiuscula P. M. Jørg. (Kenya & Republic of South Africa). Pannaria rubiginosa (Ach.) Bory (Republic of South Africa to Ethiopia, Swinscow & Krog 1988: 154). Pannaria santessonii Swinscow & Krog (Tanzania, Swinscow & Krog 1988: 155). Pannaria squamulosa P. M. Jørg. (Republic of South Africa). Pannaria tavaresii P. M. Jørg. (Republic of South Africa to Ethiopia, Krog 2000). Parmeliella coelestina Zahlbr. (Republic of South Africa, Jørgensen 1998). Parmeliella dactylifera P. M. Jørg. (Republic of South Africa). Parmeliella furfuracea P. M. Jørg. (Republic of South Africa, Jørgensen 2001c).
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Parmeliella imbricatula (Müll. Arg.) P. M. Jørg. (Republic of South Africa). Parmeliella lacerata P. M. Jørg. (Republic of South Africa, Jørgensen 1998). Parmeliella mariana (Fr.) P. M. Jørg. & D. J. Galloway (East Africa, Swinscow & Krog 1988: 154; Cameroon). Parmeliella stylophora Vain. (Tanzania, Swinscow & Krog 1988: 156; Cameroon). Parmeliella triptophylloides P. M. Jørg. (Kenya). Protopannaria pezizoides (Weber) S. Ekman & P. M. Jørg. (Algeria & Uganda, Jørgensen 1978: 53). Psoroma asperellum Nyl. (Republic of South Africa, Nylander 1863: 24). Psoroma fruticulosum P. James & Henssen (Republic of South Africa). There are additional taxa on some of the islands [Macaronesia (Hafellner 1995), Madagascar, Seychelles, The Mascarenes & Prince Edward Islands (Jørgensen 2000), St Helena, Ascension & Tristan da Cunha (Jørgensen 1977)], which are not included in the list. Discussion The Pannariaceae flora of the mountains in NW Africa (The Atlas and connected chains) is closely linked to that of Macaronesia and SW Europe, but is poorer in taxa. A Mediterranean element can also be traced in other parts of the continent. Well-known is the connection with the distant Cape Province (Almborn 1966), to which the presence of Moelleropsis nebulosa is an addition. More remarkable is the presence of Fuscopannaria ignobilis in East Africa, a region not particularly well-known for its Mediterranean elements, though flowering plants are known to have penetrated south through Egypt and Sudan, and this appears to be a possible route for the taxa reaching as far south as the Republic of South Africa. The East African mountains also contain an arctic-alpine element, for example Fuscopannaria praetermissa, Pannaria hookeri and Protopannaria pezizoides, all at high altitudes (d4000 m). Not unexpectedly, East Africa with its high mountains and varied habitats (Swinscow & Krog 1988) has the richest
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THE LICHENOLOGIST
Pannariaceae biota, ranging from the Republic of South Africa to Ethiopia, mostly containing pan(sub)tropical species, a few with Indian connections (Erioderma meiocarpum, Kroswia crystallifera), and some species otherwise restricted to South America (e.g. Erioderma verruculosum) or Australasia (Fuscopannaria subimmixta, Pannaria centrifuga and Parmeliella furfuracea)—the last three in the Republic of South Africa (for further discussion of this element, see Jørgensen 2002b). There is also a small subantarctic element in the mountains of South Africa (Psoroma), which have a varied pannariaceous flora with additional afroalpine, Mediterranean, and subtropical elements, as well as possible endemic taxa. The scarcity of the Pannariaceae in West Africa (excluding Macaronesia) is remarkable, and possibly caused by the scarce collecting in these parts of the continent. In general, Africa has remarkably few species in this lichen family compared with Australasia and South America, which appear to contain the evolutionary centres for several of the genera. The African species, on the other hand, give, with few exceptions, the impression of being remnants of once more widespread taxa. R Almborn, O. (1966) Revision of some lichen genera in South Africa. Botaniska Notiser 119: 73–112. Arvidsson, L. (1982) A monograph of the lichen genus Coccocarpia. Opera Botanica 67: 1–96. Dodge, C. W. (1964) Some lichens of Tropical Africa IV. Beiheft Nova Hedwigia 12: 1–225. Frisch, A. (1999) Afroalpine macrolichens of Mount Kenya. Bayreuther Forum Ökologie 64: 65–102. Frisch, A. & Hertel, H. (1998) Flora of macrolichens in the alpine and subalpine zones of Mount Kenya (Kenya). Sauteria 9: 363–370. Hafellner, J. (1995) A new checklist of lichens and lichenicolous fungi of insular Laurimacaronesia, including a lichenological bibliography of the area. Fritschiana 5: 1–132. Henssen, A. M., Renner, B., Marton, K., James, P. W. & Galloway, D. J. (1983) Studies in the lichen genus Psoroma 2. Psoroma fruticolosum and P. rubromarginatum. Mycotaxon 18: 29–48. Iversen, S. T. (1991) The Usambara mountains, NE Tanzania. Phytogeography of vascular plant flora. Symbolae Botanicae Upsalienses 29(3): 1–234.
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Jørgensen, P. M. (1977) Foliose and fruiticose lichens from Tristan da Cunha. Skrifter fra Det norske Videnskapsakademi i Oslo, Matematisk naturvitenskapelig klasse, n.ser. 36: 1–40. Jørgensen, P. M. (1978) The lichen family Pannariaceae in Europe. Opera Botanica 45: 1–123. Jørgensen, P. M. (1998) Studies in the lichen family Pannariaceae VII. On some poorly known Parmeliella-like species from the Southern Hemisphere. Lichenologist 30: 533–541. Jørgensen, P. M. (2000) Studies in the lichen family Pannariaceae IX. A revision of Pannaria subg. Chryopannaria. Nova Hedwigia 71: 405–414. Jørgensen, P. M. (2001a) Survey of the lichen family Pannariaceae on the American continent, North of Mexico. Bryologist 103: 670–704. Jørgensen, P. M. (2001b) New species and records of the lichen family Pannariaceae from Australia. Bibliotheca Lichenologica 78: 109–139. Jørgensen, P. M. (2001c) Four new Asian species in the lichen genus Pannaria. Lichenologist 33: 297–302. Jørgensen, P. M. (2001d) On the status of the Erioderma names in Zahlbruckner’s Catalogus. Taxon 55: 525–541. Jørgensen, P. M. (2002a) Two species of the Pannariaceae (lichenized ascomycetes) from New Zealand with a wider Southern Hemisphere distribution. New Zealand Journal of Botany 40: 327–329. Jørgensen, P. M. (2002b) Kroswia, a new genus in the Pannariaceae (lichenized ascomycetes). Lichenologist 34: 297–303. Jørgensen, P. M. & Arvidsson, L. (2001) The sorediate species of the lichen genus Erioderma Fée. Nova Hedwigia 73: 497–512. Jørgensen, P. M. & Galloway, D. J. (1992) Pannariaceae. Flora of Australia 54: 246–293. Jørgensen, P. M. & Kashiwadani. (2001) New or misunderstood species of Japanese Pannaria (Lichenes). Journal of Japanese Botany 76: 1–10. Krog, H. (2000) Corticolous macrolichens of low montane rainforests and moist woodlands of eastern Tanzania. Sommerfeltia 28: 1–75. Nylander, W. (1863) Synopsis Lichenum 2. Paris. Øvstedal, D. O. & Lewis Smith, R. (2001) Lichens of Antarctica and South Georgia – A Guide to Their Identification and Ecology. Cambridge: Cambridge University Press. Rogers, R. W. (1992) Lichen ecology and biogeography. Flora of Australia 54: 30–42. Rogers, R. W. & Stevens, N. (1981) Lichens. In Ecological Biogeography of Australia (P. L. Keast, ed.). The Hague: Junk. Stizenberger, E. (1890) Lichenaea africana. Fasc.1. Berichte der Tätigkeit der St. Gallen Naturwissenschaftlichen Gesellschaft 1889/1890: 1–144. Swinscow, T. D. V. & Krog, H. (1988) Macrolichens of East Africa. London: British Museum (Natural History).
Accepted for publication 4 October 2002