The lichens of Marion and Prince Edward Islands

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Sou/hAlnc91l Journal 01 Botany 2001 . 67 552- 572 Pnnted III South Afne;) - Alt ngl1ls reserved

SOUT H AFRICAr'OO URNAL OF BOTANY ISSN 0254- 6299

The lichens of Marion and Prince Edward Islands DO 0vstedaP and NJM Gremmen 2* , Botanical Institute. AI/egaten 41 . N-5007 Bergen. Norway , Bureau Data Analyse Ecologie. Hesselsslraal 11. 7981 CD Diever, The Netherlands .. Corresponding author. e-mail: [email protected]

Received 7 June 2000. accepted in revised form 18 February 200 1

A total of 100 lichen speci es are listed for the islands. Six species are described as new to science : Arctomia subantarctica 0vstedal , Arthothelium evanescens 0vstedal, Coccotrema stelfata 0vstedal, Ramonia subantarctica 0vstedal , The/ocarpon subantarcticum 0vstedal and Verrucaria umbilicata 0vstedal. Four new combinations are made: Amandinea subplicata (Nyl.) 0vstedal , Poeltinula cerebrinel/a (Nyl.) 0vstedal , Verrucaria nigra (Mull. Arg.) 0vstedal (from Cape Horn) and Verrucaria tesselata (Dodge) 0vstedal (from Heard Island). One third of th e islands ' lichen species also

occur in the Northern Hemisphere. Of the Southern Hemisphere species, 24 are exclusively known from areas to the east of the islands (Kerguelen Province Islands, Australia, New Zealand , contrasting to only six species recorded exclusively to the west (South Georgia, Antarctic Peninsula) . This suggests a distribution against the present predominantly westerly winds. The large number of seemingly endemic species is suggested to indicate the incomplete knowledge of the lichen floras of th e sub·Antarcti c islands.

Introduction

The lichen flo ra of sub-Antarctic Marion an d Prince Edward Islands has never been thorough ly studied. Although some accounts of the lichens of these islands have been published, these have been based on collections by non-specialists (e. g. Lindsay 1976. 1977) . or cove r only a limited taxonomical group (e.g. Hertel 1984. Henssen 1986. Henssen and Lumbsch 1985). The first account of lichens from Marion Island, based on a visit of a few hours. listed five species (Crombie 1878). The first extensive account of th e lichen flora of the islands was published by Lindsay (1977) . based on the collections of the 1965/66 Biological and Geological Expedi tion (Van Zinderen Bakker et a/. 1971). bringing the number of identified species up to 45. Further lichenological work was done in 1982 by Hertel and Henssen (Hertel 1982. 1984. Henssen 1985b. 1986. Henssen and Lumbsch 1985. J0rgensen 2000). Field observations suggested that the above accounts did not cover the whole of the lichen flora of the islands. Therefore further collections were made by NJM Gremmen in 1993/94 and 1998. The present account is mostly based on the latter collection. In ad dition a small bu t interesting collection made in 1982 by Dr A Henssen was placed at our disposal. A few species thai have not been fou nd in the above-men tioned collections but were recently published and are available in herbaria have been included . Unfortu nately the collection on which lindsay's paper was based , which was housed in Bloemfontein , could not be traced (see Discussion).

The microscopical and ch emica l studies followed conventional me thods.

The islands Marion and Prince Edward Islands (4rS. 38'E) are situated in the Southern Ocea n. some 2 OOOkm southeast of the southernmost point of Africa. just north of the Antarctic Polar Front. Together these island s make up the Pri nce Edward Islands group. one of the few island groups in the Southern Ocean . The nearest land is the Crozet arc1ipelago, some 950km to th e east. Marion Island measures 290km2 , and reaches an elevation of 1 230m. It is 22km removed from Prince Edward Island. which is 90km'. with its highest point at 672m. The islands represen t the summits of shield volcanoes, risi ng from the West Indian Ocean Ridge. (Verwoerd 1971). and are geologically relatively young. There appear to have been different periods of volcanic acitivity, and two distinct series of basaltic lava can be discerned. The older series consists of massive grey lavas dated at about 275 000 to 100 000 years (McDougall 1971). The younger series of scoriaceous, blocky black lava is not older than 15 000 years. The youngest lava flow dates from 1980. Much of the land surface consists of lava, and of scoriae deposits. In the lowlands immature soils have developed (Smith 1976). Larg e tracts of th e lowland areas are covered by mires, in which grasses and bryophytes are dominant, and which have given to rise to deep peat deposits.

South African Journal of Botany 2001 , 67 . 552-572

The islands have an extremely oceanic climate, with low temperatures (average 5°C) , showing liltle diurnal (5°C) or seasonal (4°C) variation (Schulze 1971). At sea level frost occurs some 55 days on average per year. Precipitation exceeds 2 500mm. yr1 and is mostly in the form of rain, although snow may fall in any season . Winds are usually strong , reaching gale force on over 100 days per year, and are predominantly from the west. Humidity is high (> 80% average), as is cloudiness, and sunshine reaches only about 30% of the maximum possible (Schulze 1971 ), During the past 300 000 years Marion Island has been subjected to three glacial episodes, the last of which ended ca, 12 000 years ago, during which temperatuers were some 3.5°C lower than at present (Hall 1978, Van Zinderen Bakker 1973). However, there is no evidence for glaciation on Prince Edward island. Recent decades have shown an increase in the mean annual air temperature of ± 1°C (Smith 1992), and a significant decrease in precipitation (Chown and Smith 1993), The island's biota reflects its strong oceanic cha ra cter: the number of species is low (e.g. 24 indigenous vascular plants, some 80 mosses, and 45 liverworts), and some species groups which playa major role in many conti nental ecosystems (e.g. land mammals, herbivores as well as predators) are absent in the native biota. The islands are breeding grounds for large num bers of seabird s and seals, locally resulting in heavy nutrient enrichment. A number of alien plants and animals have been introduced , e.g. cats, mice, invertebrates and vascular plants (Gremmen 1982, Gremmen and Smith 1999, Watkins and Cooper 1986). The terrestrial vegetation of the island can be classified into six comm unity complexes (Gremmen 1981 , Table 1), The distribution of the plant communities is largely determined by differences in moisture reg ime and nutrient status be tween the different habitats (Gremmen 1981 , Smith 1987), In additio n there are communities of epilithic habitats, No detailed information on the species composition of these

553

communities is available. The main distinction is between coastal areas , influenced by sea wate r, where T urgidosculum complicatulum, Verrucaria sp. and Caloplaca sp, are abundant (Fuller 1967, De Villiers 1976), and inland epilithic habitats. Communities of crustose lichens are common on many rock surfaces, but commu nities dominated by macrolichens, e.g. Usnea or Stereocaulon, cover only very small areas, The general distribution of habitat types is illustrated in Figure 1, The species Acarospora cf, otagensis H, Magn, Thallus as elongate, convex lobes, ca, 3mm long and 0,5mm broad, pale grey-brown, Apothecia sunk in thallus, disc red -brown, ca, 0,3mm diameter. Spores elliptic, ca, 4 x 1,5mm, TLC: not investigated due to small material. Ecology: on coastal cliffs, with Amandinea subplicata and Caloplaca spp. Distribution: New Zealand, Marion lsI. Comments: the material is very sparse, but it com es close to the description in Galloway (1985) except that the New Zealand material has an areal ate thallus, The type has not been available. Material seen: Gremmen 94- 269, Marion Island, Trypot Beach, 4m a.s.I. , black la va rock outcrop in area with seals and penguins; 30m from shore, saltspray, slightly sheltered, Amandinea subplicata (Nyl.) IZlvstedal comb , nov, Basionym: Lecidea subplicata Nyl. , Ny!. in Crombie 1877, J, Linn, Soc, London , Bot. 15: 190, Thallus crustose, effuse, rimose , buff, thin, with a broad, brown pro thall us, Apothecia up to 0,5mm diameter, dark brown to black, at first sunk in thallus, later half-emerged , with excipulum proprium only; margin protrudi ng. Often cracked; disc flat, often with a reddish hue, and usually

Table 1: Marron Island vegetation complexes. The communities of 1-6 and 9 are described in Gremmen (1981). Communities of group 7 are partly described in De ViJliers (1976) . Communities of group 8 have not yet been described

1. 2.

3.

4. 5. 6

7. 8 9.

Crassu/a rnasc/1ata com plex vegetation of areas strongly influenced by wind-blown sea spray CalliMche antarctica-Poa cookii complex commumties of areas in nuenced by trampling and nutrient enrichment by animals (seals, penguins and other birds) Acaena mageJlanica-Brachythecium complex communi ties of areas with a pronounced la teral. mostly subsurface , water now (springs, flushes , and drainage lines). usually dominated by Acaena magellanica. with a weH developed bryophyte stratum. Juncus scheuchzerioldes-Blepharidophyllum densifo/ium complex mire vegetation, on wet, nutrien t-poor peat soils, usually dominated by hepatics, mosses. and graminoids Bfechnurn penna-marina complex commu nities on mineral soil s on well-drained lowland slopes, dominated by the small fern Blechnum penna-marina Andreaea-Racomitrium crispulum complex communities of areas strongly exposed to the wind , characterized by low plant cover and the dominance of the cushion forming vascu lar plant Azorella selago and cushion forming mosses. These communities cover over 50% of the lowland areas, and dominate the vegetation above 300m altitude. communities of coastal epililhic habitats epifithic communities in areas strongly exposed 10 seawater or salt-spray communities of inland epilithic habitats epililhlC communities not in fluenced by salt-s pray or seawater aquatic communilies

0vsledal and Gremmen

554

V IV II

W

IV

III

III

m 1200

II 800

~I

0

1

3

2

4

5 km

I

rr

E

400 0

Figure 1: Schematic east-west profile of Manon Island . indicating the distribution of the genera l habitat Iypes. J = coasta l epillthic habitats (vegetation complex 7) : II = coastal habitats (vegetation complex 1 and 2) : III = lowland areas (complex 3, 4 . 5, 9, some 2 and 6), IV = upland areas (vegetation complex 6 and 8); V = central mountain area (ice, bare rock. and complex 8) . (modified after Sm ith 1987)

gyrose. Hymenium 10 0- 11 0~m high , epithecium brownish, N-. Spores 8 in asci, 10- 15 x 6-7~m . Paraphyse end cells ca 4~m wide. Conidia strongly curved , ca. 15 x l~m. TLC: negative. Ecology: on black and grey lava coastal rocks, together with Ca/oplaca lucens, Arthothelium evanescens and other lichens. Distribution: Kerguelen , Marion 151. , Prince Edward 151. Specimens seen: Gremmen 94-2 69 , 93-014, 94- 271 , 94-70,94-71,94-72,94-2 63,93-016, Henssen 28 061, 28 096b, 28 054a, 28 004e, 28 139. Comments: the specimens have been compared with the type from Ke rguelen (H) and fou nd to be similar in all essential characters. H. Mayrhofer (pers. comm.) placed this taxon in Amandinea. Arctomia subantarctica 0vstedal sp. nov. (Figure 2) Arctornia interfixa affinis, sed thalia majore, apothecia semi-

.7-. ' f:

B

A

.. .. ~

•• '

'

• ,. I

:.:;"':::<:<.::!:: 't • I

. ' '.

: '.'','.':\

..., -:

,

1 mm

.

.

.

,

;~ ,"

..

:. . 1 mn

c

1 mm Figure 2: Arctomia subantarctica. A: vertical section of part of apothecium : B: spore . Horotype

immersa et cortices numerosis stratis cellulis compositi. Holotypus: vide infra. Thallus foliose, small , lobes squamulose or elongate, forming rosettes, individ ual lobes 0.4- 0.6mm broad , 1- 2mm long, flat, brown . Lower suriace grey-brown. with a few, very short rhizinae. In transect the thallus is 270-280~m high, with an upper, paraplechtenchymateous, 5-6 cells high cortex, 20 - 25~m high, brownish; lower cortex paraplech tencymateous, 30- 35~ m high , 10-12 cells hig h, pale brownish. Medulla with strands of fungal hyphae and colo nies of Nostoc. Apothecia 0.5-0.6mm diameter, half sunk in squamules, proper margin slightly raised , paler than thallus, disc fiat , brown. Excipulum proprium composed of a paraplechtenchymateous ti ssue. Hymenium uncoloured, 60-70llm high, uppermost part pale brownish. Hypotheciu m colou rless. Paraphyses simple, end cells somewhat enlarged , with constricted septa. Asci narrowly clavate, of Bacidia type. Spores 8 in asci, uncoloured , narrowly elliptic, 5-6-septated, 28- 32 x 4-5~m. Pycnidia not seen . TLC: negative . Ecology: on epilithic bryophytes (Grimmia sp., Andreaea spp .), togethe r with Massa/ongia camosa, Siphu/astrum papillatum, Pannaria pulvinufa. , etc . Specimens seen : Gremmen no 94-245, Prince Edward Island, N slope of Kent Cra te r; 250m a.s.l.; on cus hions of Grimmia and Andreaea on boulders, 15.5. 1994 (holotypus BG); 94- 235. Comments: the specimens were at first determined as Massa/angia carnaS8, but the lack of isidia, the semiimmersed apothecia and 5-6 septated spores showed that it was something different. The other two species known in the genus, A. delicatula and A. inierfixa , have less developed thalli with cortex only one layer th ick and convex apothecia wi th excipulum proprium consisting of radiating hyphae (Henssen 1969). However, ascus type, spores and paraphyses are similar in all three species. Arthothelium evanescens 0vstedal sp. nov. (Figure 3) Thallus epilithus, effusus, rimosus, grisea-lividus. apothecia 0.3-0 . 5~m la ta, convexa , nigra, emarginata. Hypothecium subfuscum. Sporae ova tae, murales , septis horizonlalibus 6-7, septis verticalibus 3-4, decolores, 28-32~m longae, 8 -13~m latae. Holotypu s: vide infra.

555

South African Journal of Botany 2001. 67: 552- 572

Ecology: on mosses and lichens.

A

Distribution : Marion lsI.; cosmopolitan in colder areas.

Specimen seen: Henssen 28 11 4c.

LJD

Biatora sp. Thallus invisible, apothecia situated on algal matrix on

00 E

"a

M

Andreaea spp. Apothecia brown-black , convex, up to O.5mm

00 L:=J [70 l

E

"a

'"

diameter. Excipulum proprium 60- 70lJm broad , colourless

except outermost part which is red-brawn-pigmented , composed of a radiating , anastomosing hypha I web embedded in a gelatinous matrix. Hymenium 45-55~m high, colourless except uppermost part which is red-brown. Hypothecium

J

red-brown , 60-70~m high, composed of irregularly oriented hyphae. Subhymeniallissue colourless, extending deep into th e algal matrix . Asci of Biatora (?)- type, ca 25 x 8~m. Spores 8 in asci , uncoloured , 1-3-septated , 7-10 x 2 .5- 3~m . Paraphyses ca 1~m broad , flexuose , strongly

c=::7 \::7

ram ified to anastomosing, end cell not enlarged but embedded in the epithecium matrix. Figure 3: At1hotl1elium evanescens. A: spore: B: ascus with paraphysoids. Holotype

TLC: not performed. Ecology: on algal matrix on moss cushions (Andreaea sp.) in feldmark

Thallus effuse, crustose, very thin , rimose to granulose, pale

Specimen seen: Gremmen 94-214. Comments: the details of the apothecia point to the genus Bialora, as defined by Printzen (1995). In the European

Distribution: Marion Island.

grey to brown-grey. Apothecia sessite , emarginate, btack, 0.3-0.4mm in diameter, 150-200~m high , in outline roundish but often somewhat irregular, slightly convex , sur-

flora , there are two species which have somewhat similar spores: viz. B. aegrefaciens Printzen and B. rufidula

face rough . Hymenium 60 -70~m high, K/ t+ wine-red, epith ecium wi th btackish granules. Hypothecium dark brown. Asci semiglobose, ca 50~m long, KlI+. Spores 8 in'

(Graewe in Hellb.) S. Ekm . and Printzen (Printzen 1995) but both species have larger spores, a distinct thallus and are growing on bark.

asc i. muriform, uncoloured , with 6-7 transverse and 3-4

vertical sepia , 29- 32 x

8-13~m.

Epispore very thick .

Buellia fuscoatratula Zahlbr.

Pycnidia not seen.

Thallus crustose, effuse, strongly rimose , rusty brown , mar-

TLC: three unknown compounds (J.A. Elix teste). Ecology: in splash zone on coastal rocks , 2m a.s.l. , with Amandinea subplicata, Caloplaca spp., Turgidosculum complicalulum .

ginal lobes not clearly differentiated . Apothecia abundant, half sunk in areolae, up to 0.5mm diameter, black, proper margin slightly raised, somewhat paler than disc, disc flat. Hymenium 70 -80~m high , epithecium brownish, N+ red . Spores 8 in asci, brown, 14-18 x 7-10~m , septum constricted, wall minutely warted. Paraphyse end cell enlarged,

Specimens seen:

Gremmen

94-263 , Marion

Island ,

Macaroni Bay (holotypus BG), 93-014 , 93- 016. Comments: It has not been possible to identify this species with any saxicolous Arthothelium species included in

Redinger (1937), nor with any known from Europe (Poe It 1969 , Purvis et al. 1992) or New Zealand (Galloway 1985). On the coastal rocks of the Chilean Pacific coast, three Arthothelium species occur (Follman 1968). Two of these , A. halophilum Follm. and A. pacificum Follm., have immersed apothecia and wel l developed thalli and are clearly different. The third , A. spilomatoides (Ny!.) Zahlbr., has sessile apothecia and a less well developed thallus. Some Chilean

ca, 41.1m diameter. Pycnidia not seen.

TLC: unidentified xanthone. Ecology: close to stream on the plateau, on rock , with Ca/opolaca sublobutata. Distribution : New Zealand (Galloway 1985), Marion Island. Specimen seen: Henssen 28174.

Comments: the specimen has been compared to the type (W) and is fairly similar in most essential characters, but differs in somewhat larger spores (10- 14~m long in type) .

specimens (8) were examined and were found to have a

Caloplaca castellana (Rasanen .) Poelt

much better developed thallus than in the present species,

No visible thallus present , apothecia sunk in cyanobacterial crust. Apothecia lecanorine, up to OAmm diameter, with pale orange-yellow disc and grey-green margin. Hymenium ca

and also distinctly smaller spores, showing it is not the same species .

Arthrorhaphis citrinella (Ach.) Poelt Thallus crustose , greenish yellow, areolate , areolae often dispersed, to O.5mm diameter, sorediate at marg ins, without a epinecrallayer. Apothecia not seen . TLC : rhizocarpic acid.

70~m high , excipulum proprium ca 15mm wide, composed of short, broad cells. Excipulum thallinum ca. 100~m broad, cortex weakly developed, inner part almost completely filled with algae, which extend below hymenium. Spores 8 in asci, 11-12 x 5-6, septum ca 6~m. Paraphyse end cell slighlly enlarged to 2-2.5~m , epipsamma little to medium coarse.

0vsledal and Gremmen

556

Oil paraphyses not seen. Pycnidia not seen.

TLC: not performed. Ecology: on lower surface of flat stone in supralittoral , with

Amandinea subplicata , undetermined sorediate, Turgidosculum complicalulum. Distribution: Europe, N. America , Marion Island. Specimen seen: Gremmen 94-265 pp.

and

Ca/op/aca cirroclrrooides (Vain.) Zahtbr. Thallus placaid, as small rosettes, 4-5mm wide , with ca O.2mm wide , convex, radiating lobes, orange-yellow, sorediate. So ra lia tip-shaped, developing from th e lobe-tips. Apothecia not seen.

TLC : not performed. Ecology: on coastal rocks, just above intertidal zone. Distribution: Marion 151.; nearly circumaustral. Specimen seen: Gremmen 94-265 pp. Ca/op/aca cribrosa (Hue) Zahlbr. Thallus 1-2cm diameter, lobate, sulphur yellow, lobes thick,

Calop/aca /ucens (Nyl.) Zahlbr. Thallus placoid, up to 4-5cm diameter, lobate at margins . Lo bes discre te, narrow, ca. 1 mm wide, co nvex, orange.

Central part with papillae, ca 0.5 x 1mm, often wi th pseudocyp hellae on top . Apothecia leca norine, sessile, up to 1mm

diameter. Hymenlum 100-110 ~ m high, with fine epipsamma. Spores 8 in asci, 14-15 x 6-8~m , septum 5.5-6~m. Oil paraphyses present. TLC: not performed . Ecology: mostly on coasta l rocks , with Ca/op/aca sub/obu/ata, Lithographa spp., etc. , but occasionally up to 150m above sea level. Distribut ion: Marion lsi., Kerguelen , Antarctica.

Specimens seen: Gremmen 94-47, 93-032, 93-015, 94-43, 94-265, 93-003, 94-269, 94-41, 94- 262, 93- 106, Henssen 28003a, 28010a, 28004a 28068. Com men is: there seems to be no differences from western Antarctic populations.

radiating at margin, in inner part vertical. with slit-like

Ca/op/aca sub/obu/ata (Nyl.) Zahlbr.

pseudocyphellae. Apothecia not seen. Ecology: in intertidal and on coastal cliffs, wi th in the saltspray zone; with Amandinea subp/icata and Arfhothetium

Thallus either crustose or effigurate, up to seve ral cm in diameter, yellow orange to orange. Apothecia with double margin, sessile , up to 1.5mm diameter. Disc when young plane, becoming convex, dark orange. Margin prominent when young, later becoming much thi nner. True margin somewhat lighter orange than the disc, thalli ne margin even

evanescens. Distribution: Marion Island , Australia, and New Zealand , Incl uding Auckland and Antipodes Islands (Poel t and Pelleter 1984). Specimens seen : Gremmen 94-2 63, 94- 269, Henssen 28010 b. Calop/aca isidioc/ada Zahlbr. Thallus composed of finger-like reddish orange isidia , sometimes organised into rosettes with flattened marginal lobes. Apothecia rare , lecanorine to zeorine . Spores 8 in asci,

12-16 x

6 -8~m,

septum

5-6~m.

Paraphyse end cell ca

4~m.

Ecology: on lichens on rocks at low and high elevations. Distribution: Marion lsI., Juan Fernandez, Tristan da Cunha,

South Georgia, South Orkney 151., Antarctic Peninsula, Ke rguelen. Specimens seen : Gremmen 94-72 , 94- 73, Henssen 28140c, 28055, 28137 a. Comments: The C. isidiocJada materia l comes from a large area, and varies in dimensions of isidial outgrowths, apothe~

cium type, spore form , colour, and host (Table 2). Whether these are genotypic differences, is not yet clear. The varia-

tio n of the populalion should be studied in the field .

lighter, concolorous with tha llus. Thalline exci pulum almost filled with algae, and with a paraplechtenchymatous cortex. Proper excipulum well developed , up to 50~m thick, composed of prosoplechtenchymatous tissue. Hymenium 110-120~m high, with medium coarse epipsamma. Paraphyses ca. 1.5~m diameter, end cell enlarged to ca. 3 ~m diameter. Asci with 8 spores , spores 14-16 x 7-8 ~m , septum 3.5-5~ m. Ecology: Widespread and common on maritime cliffs , rocks and stones, sometimes weI! away from the direct influence of the sea but usually close to bird colonies; ornithoco~

prophilous. Distribution: Marion lsI., Prince Edward 151. , South Africa, South Australia, New Zealand , southern Chile, Tierra del Fuego, Staten Island , Falkland Is., Tristan da Cunha. Antarctic: South Georgia, Bouvel0ya , South Sandwich Is., South Orkney Is .. South Shetland Is., Anta rctic Peninsula. Specimens seen: Gremmen 93-014, 94-47 , 94-269, 93-016, Henssen 28058b Comments: listed by Lindsa y (1971) as C. depauperata (MuII.Arg.) Zahlbr.

Table 2: Some characlers of various populations of CaJop/aca is;dloc/ada coli. Width of isidial outgrowths

Spore sIze

Wes tern Antarcl1c area Manon and Prince Edward Islands Kerguelen

100IJm

"m 12- 14 x 6 5-8

45IJm

12-16 x 6- 8

36IJm

13- 14 x 7-10

Tristan da Cunha

28IJm

?

Spore form

Apothecium margin

Narrow elliptIC

Lecideine

Pyriform to Lecldeme (rarely broadly elliptiC zeorlne) ElliptiC to Lecideme (rarely broadly elliptiC zeonne) ? ?

Oil paraphyses

Disc colour

Ves

Concolourous with lhallus Concolourous with thallus Darker than thallus

No no ?

?

Substrate/host MaInly on base of fruticose lichens On CfUsLose lichens On crustose lichens On cyanobacteria

SOUlh Afocan Journal of Botany 200 1. 67 552-572

Caloplaca tiroliensis Zahlbr. Thallus invisi ble or as a very thin , pale ochraceous crust . Apothecia up to 0.5mm diameter. sessile. thin. yellow to pale orange-yellow; proper margin thin. slightly protruding. slightly paler than disc; disc flat to slightly convex, smooth. Thalline margin mostly not present. Hymenium 60-70~m high , epipsamma medium coarse. Spores 8 in asci , 12-14 x 6 - 7~m, septum ca 4~m. Paraphyse end ce ll ca 3~m across. TLC: not performed. Ecology: on crustose lichens, in one case on bryophytes . Distribution: Marion lsI. , cosmopolitan in cold areas. Specimens seen ' Henssen 28114c, 28 114d. 28174a. Comments: somewhat different from western Antarctic poputations (0vstedal and Smith 2001): smaller spores, narrower paraphyse end cells, mainly tecideine apothecia , but within the varialion of European popu tat ions (see Poelt 1969).

Carbonea assentiens (Nyl.) Hertel Thallus crustose, effuse , thin , strongly rimose , wi th slightly bullate areoles, grey with a yellowish tinge. Apothecia ses sile. fla t, up to 1mm diame ter. black , with proper margin slightly protruding. Hymenium 35-40~m high , epithecium green-brown. Excipulum propnum and hypothecium dark brown to black. Spores 8 in asci, simple, uncoloured , 9-11 x 4 -6~m.

TLC : atranorin , zeorin , unidentified xanthone. Ecology: on rock at low to medium elevation, with Lecanora disjungenda. and other lichens. Distribution: Ma rion lsi" Prince Edward Island, Kerguelen , Antarctic (see , however, comments below) Specimen seen: Gremmen 94-73. Commen ts: C. assentiens s.str. , described from Kerguelen, appears to be restricted to Kerguelen and Prince Edward Islands. In the western An tarctic region. there occu rs a closely related taxon which has been included in the same species (Hertet 1984). It has a thicker. white thallus with no xanthone , higher hymenium and larger spores.

Carbonea phaeostoma (Nyl.) Hertel Thallus crus tose. effuse , very thin, grey, strong ly rimose. Apothecia sessile, up to 1.7mm diameter. proper margin black. strongly protruding. disc flat , ci nnamon brown. Hymenium 80-9 0~m high , epithecium red-brown . containing crystals of an thraquinone 2-chloremodin (Hertel 1984) . Excipulum proprium and hypothecium black. Spores 8 in asci, simple, uncolou red, 11-13 x 5-6 ~ m. TLC: atranorin , zeorin, 2-chloremodin. Ecology: on coastal rocks , with Calillaria impolita, and other lichens; also inland from low to high elevations. Distribution : Marion Island, Prince Edward Island, Kerguelen. Specimen seen : Gremmen 94-135. Carbonea superjecla (Nyl.) Hertel Thallus within host. Apothecia black, 0.3-0.5mm diameter, half sunk in thallus to sessile. true margin broad , fissured , disc with umbo. Hypothecium and excipulum brown-black. Hymenium 50 -80~m. Spores 8 in asci , ellipSOid , 10-15 x 5- 8~m. Paraphyse end cells en larged to 3.5~m diameter. Ecology para symbion t on crustose lichens .

557

Distributio n: Prince Ed ward Isla nds. Kerg uelen Specimen not seen ; distributed in Lecideaceae Exciccatae (Hertel 1982). Catillaria impolita D.C. Lindsay Thallus crustose. effuse, very thin dark grey. Apothecla sessile. up to 1.2mm diameter, black, excipulum thallinum absent, excipulum proprium protruding. Hymen ium 50-70~m high . red-brown in thick sections, epithecium green-black. Excipulum and hypothecium dark red-brown. Spores 8 in asci . uncoloured , 1-septate. 11 - 13 x 5-6[Jm. Paraphyses with end cells enlarged to 2-4~m diameter, constricted at septa. TLC: negative. Ecology: on black lava rocks in lowland area, near coast. Distribution: Marion , Prince Edward lsI. Specimen seen: Gremmen 94-135. Comments: Lindsay (1976) described C impolita as new from Marion Island and the prese nt speci men conforms well with the description , except that the spores are smaller (18 x 8 ~ m in Lindsay I.c.). The presen t specimen cou ld also be included in C. chalybeia, a common species on sea shores in N Europe, except that the paraphyse end cells are larger in that species. More material is necessary to solve the relationships. The type of C. impolita has not been available.

Cati/laria lenlicularis (Ach.) Th. Fr. Thallus very thin , effuse, brown-grey. Apothecia sessile to half immersed, up to O.2mm diameter, brown, proper margin distinct, when wetted appearing distinctly darker than disc. Hymenium ca. 30~m high. Excipulum proprium golden brown . composed of an intricate we b, with somewhat radiating hyphae. Asci ca. 20 x 1O~m, with a th ick tholus . Spores 8 in asci. uncoloured , 1-septated . ca 8 x 3~m. Paraphyses mos tly simple, with distinct enta rged end cell with brown cap. Hypothecium golden brown. TLC: negative. Ecology: on rock, close to stream on the plateau, with Buellia fuscoalratula and Caloplcea cf. sublobulala. Distribution: Marion lsI. , Europe. Specimen seen: Hensssen 28 174. Cladonia albala Stenroos Primary thall us squamulose, squamules up to 3mm long . Podetia up to 2cm high , green-brown, simple to branched, scyphi branched by margins proliferating, always scyphose. Scyphi up to 7mm wide, graduatly flaring. Podetia surface corticated; inside scyphi white, showy granules. Hymenial discs brown. TLC: fumarprotocetraric acid complex. Ecology; on soil. Distribu tion: Prince Edwa rd Island endemic Specimen not seen , description based on Stemoos (1993). Cladonia celVicomis (Ach.) Flot. ssp. mawsonii (Dodge) Stenroos and Ah ti Primary thallus squamulose, squamules up to 2mm long. Podetia up to 4cm long, 1-2mm broad, blackish at base, with centrally proliferating scyphae . Scyphi up to 4mm broad, closed . Surface of podetia rough ty areolated, with

0vsLeda i and Gremmen

558

microsquamu les . Hymenial disc brown. TLC: fumarprotocetraric acid complex. Ecology: on soil. Distribution: South Orkney lsI. , Prince Edward

Specimen not seen, description taken from Stenroos (1993).

lsI.,

Macquarie lsI.

Specimen not seen: description ta ken from Stenroos (1993). Cladoma chloro(Jhaea (Florke ex Sommerf.) Sprengel Primary tha llus as densely crowded ascending lobes, 2-3 x 0.5-1mm , brown. Podetia 7-10mm high , usually with relative ly broad scyphi , 3-4mm broad in old speci mens, brown-

grey, cove red with small granules and lobules. TLC: Fumarprotocetraric and protocetraric acids, Cph-1 , Cph -2. Ecology : On soil. Distribution : Marion 151. , Cosmopolitan . Specimen not seen , description taken from Stenroos (1993).

Cladonia marionii Stenroos Pri mary thallus squamulose, squamules brown-green , up to 3mm long . Podetia up to 3cm high, wi th out scyphi , whitegreen, moderately branched in upper part, axils closed to open. Podetia surface with coarse granules and microsquamules. TLC: fumarprotocetraric acid complex. Ecology: on soil deposits in open fernbrake Distribution : Marion Isla nd endemic. Specimen seen : Gremmen 94-013. Cladonia ochrochlora Florke Primary thallus squamulose, squamu les up to 3mm long . Podetia up to 3cm high, 2mm thick, green-brown, simple or bran ched, subulate or with scyphi. Scyphi up to 1.5mm wide, shallow. inside corticate. Podetia surface col"ticate and squamulose a t base , patchily farinose soredia te towa rds apices.

Hymenial disc brown. TLC: fumarprotocetra ric acid complex . Distribu tion: Marion [sl. , cosmopOlitan. Specimens not seen, description taken from Stenroos (1993).

C/adonia pocillum (Ach .) O.-J . Rich. Primary thallus warm brown , appressed to subs trate, up to 3cm diameter, individual lobes usually litt le differen tiated , when differentiated ca . 3 x 2mm . Pod etia up to 15mm high, upper part microsquamulose, lower part verruculose to corticate , scyphi up to 9mm wide, closed. TLC: Fuma rp rotocetraric acid complex. Ecology: on soil in lowland feldmark Distribution: Marion and Prince Edward [sl. , cos mopoli tan . Specimen seen: Gremmen 93-11 2. C/adonia cf. poeciloc/ada Abbayes Primary tha llus insignificant. Podetia up to 4cm high , greenbrown, simple to branched , with scyphi. Scyphi up to 3mm wide, deep, perforated, wi th marginal proliferation s. Podetial surface corticate, with aggrega tions of granules in upper part. Hymenial disc brown. TLC: fuma rprotacetraric acid complex. Ecology: On soil. Distribu tion: Prince Edward lsI., cosmopolitan.

C/adonia sarmentosa (Hook. and Tayl.) Dodge . Primary thallus squamulose, squamules up to 2mm long . Podetia up to 3cm high , 1- 1.5mm wide , brown-grey, subulate or with scyphi. Scyphi 1-1.5cm wide, closed. Podetium surface with granules and squamules. Hymenia[ disc brown. TLC: fumarprotocetraric acid complex, ± atranorin. Ecology: on soil. Distribution: Australi a, Tasmania , New Zealand , Campbell lsI. , Marion [51. , Southern South America , Antarctica . Specimens not seen, description based on Stenroos (1993). Cladonia cf. tapperi Ahti and Krog Primary thallus squamulose, squamules erect, up to 2mm long , Podetia up to 1.5cm high , without scyphi , 1-2mm th ick, green-brown, sometimes branched. Pod etia surface cortica te at base, becoming microsquamulose towards apices. TLC: fumarprotocetraric acid complex, stictic, cons tictic. Ecology: on gravelly soil (scoriae) . Distribu tion: E Africa, Prince Edward 151. , Marion 151. Specimen seen: Gremmen 94 - 98. Coccotrema stellata 0vs tedal sp. nov. (Figure 4) Thallus crustaceus, roseus , arelatus, areolae dispersa tae , saxi cola: sorediis isidiisque absentibu s. Ascocarpi a typica per genum , sed ostiolis stellatis et basis non constrictis. Sporae monostichae, 15-18 x 9-1 1 ~m , 8:nae, decolores late ellipticae. Pycnocon idia non vi di. Phycobio nt Chlorococcal es . Cephalodia irregulariter hemispheri ca, 0.5-1 .5mm diametro, algam Ca/otrix sp. co ntinens. Thallus acidum sticticum continens. Holotypus: vide infra . Th allus crustose, pink, several cm in extension, up to ca 1mm thick, weakly rimose. Apothecia l warts surrounded by cracks, up to 1.3mm in diame ter. Apothecia sunk in warts, warts with ta pering margins, ostiole irregularly stellate. Inner cavity 340-370 ~m wide, hymenium colou rless , 1 60- 17 0~m high . Excipul um proprium colourless, composed of an intricate hyphae web, 30-4 0~m broad . Asci cylindrical, ca 100 x 12~m , with a thin , lightly amyloid tholus . Spores Simple, colourl ess, with double wa ll, broadly ell iptical, un iseriate,

Figure 4: Coccotrema stellata. Scale = 1mm. Holotype

South African Journal of Botany 2001 . 67 552-572

559

14- 17 X 9-11~m. Paraphyses thin, flexuose , simple, persiste nt. Hymenial gel Kli + blue. Periphyses in lower part of ostiole cavity, thin, short, flexuose. Cephalodia up to 2mm diameter, flattened semiglobose, pale ochre, rad ially fissured, with Calotrix sp. (?). Pycnidia not seen . TLC: Stictic acid complex.

Thallus crustose , effuse , thin , green-grey; photobiont Pseudotrebouxia . Apothecia mostly sessile, up to 0.5mm diameter, brown-black ci rcular. Hymenium 95-150~m high, strongly gelatinous, brown-black in uppermost part.

Ecology: on lava rock at medium elevations.

muriform, at maturity cells transforming into co nidia .

Distribution: Marion Island, N of Junior's Kop; 160m a.s.l. ; on

TLC: not performed.

rocks in tran sitional zo ne between feldmark and flush area ,

Ecology: on rock in feldma rk at medium elevation Distri bution: Marion and Prince Edward Island, endemic.

locally common. Gremmen 94-184; 14.5.1994 (holotype BG). Oth er specimens seen: Gremmen 94-185. Commen ts: no similar species is found in Brodo (1972),Galloway (1985) or Purvis et a/.(1992). Dimerella sp. Thallus very thin, cru stose, green-brown, wi th Trentepohlia.

Apothecia almost urceolate, up to 0.3mm wide and high , flesh-coloured , disc concave , pale orange. Excipulum proprlum composed of a pseudoparenchymateous tissue , with a few algae included at margin. Spores 8 in asci, co lourless ,

1-septated , constricted at septum, ca 10 x 2~m , with epispore. Paraphyses unbranched, constricted at septa, end cell not distinctly enlarged. Pycnidia not seen. TLC : not performed . Ecology: in pits in lava rocks at medium el evation, together

with P/acopsis macrophtha/ma. Distribution: Prince Edward Island .

Specimen seen: Gremmen 94-131 . Edwardiella mirabilis Henssen Proper thallus not present. somatic pha se consisting of

hyphae imbedded in the gelatinous matrix of colonies of G/oeocapsa and other cyanobacteria. Apothecia thallinocarpous, up to 0.15mm broad, blackish when dry, when wet swelling and becoming red-brown. Hymenium 50-70 ~m high, Kli + wi ne-red. Paraphyses branched and anastomosing. Spores 2-8 in asci. Spores colourle ss , simple, 6-8 x 4.5-6~m .

Conidia rod-shaped , ca. 1.5 x 1~m . TL C: not performed .

Ecology: on cyanobacteria colonies on rock . Distribution: Marion Island endemic.

Comments: description based on Henssen (1986) of thi s primitive lichen , wh ich according to Henssen (1986) belongs in Lichinaceae.

Paraphyses branched and anastomosing in a gelatin ou s

matrix. Spores 2-4 in asci , broadly ellipsoid , uncoloured,

Specimen seen: holotype (MB) , Gremmen 94-252. Comments : A more detailed description found in Henssen

and Lumbsch 1985. Haematomma erythromma (Nyl.) Zahlbr. Thallus yellow, effuse with variable thickness, up to 10cm diameter. usuall y rimose, corticate , sorediate. Sararia coarse, not sharply delimited . Apothecia blood red , when yo ung sunken in thalline verrucae , with a very th in pale prop ~ er marg in , when aId marg in excluded and disc convex. Hymenium 60-70~m high , colourless. Paraphyses thin , flexuose, strongly branched. Spores narrowly ellipsoid , simple to 2-septate, colourless, 17.22 x 5-5.5~m . TLC: Placodial acid, pseudoplacoidal acid, arthoth el in, atranorin chloratranorin, lichexanthone, 6~0 ~meth yla rt hoth e l i n , 2-chlorlichexanthone, 2,4-dichlornorlichexanthone, 4,5dichlornorlichxanthone, stictic acid complex. Staiger and Kalb (1995) found combina tions of these compounds as four different chemotypes. Ecology: on rock. Distribution : Tierra del Fuego, Falkland lsI., Marion lsI., South Georgia , South Orkney lsI., South Shetland lSI. , Antarctic Peninsula (to ca 68°S) . Com ments: not present in our material , but reported in Staiger and Kalb (1995). Herle/fa subantarctica Henssen Th allus filamentous , olive brown , consisting of Scytonema

threads, 8 - 12~ m broad, surrounded by a gelatinous sheath in which fungal hyphae extend; altogether making up tufts ca 20mm broad and 1mm high . Apothecia olive-brown to blackish , up to 0.3mm broad, at first flat with a pale excipulum proprium , later convex with margin excluded. Hymenium 5070~m high , olive brown in uppermost part; hypothecium redbrown. Excipulum proprium composed of pseudoparen ch y-

mateous tissue . Paraphyses simple or branched in upper

Fuscidea asbolodes (Nyl.) Hertel and V. Wirth Thallus crustose , rimose, th ick, grey, up to ca. 1cm diameter; margin distinct (almost subeffigurate), blackish . Apothecia crowded in inner part of tha llus, often angular, dark brown , up to 0.5mm diameter, margin thick and protruding , disc concave. Hymenium 70 - 80~m high, upper part pale brown. Excipulum proprium black. Hypothecium pale. Paraphyses thin, flexuose , simple. Spores 8 in asci, simple, without halo, pale brown when old, 7-12 x 5-7~m . TLC: negative. Ecology: on rock from low to medium elevations Distribution: Kergu elen , Marion Island . Prince Edward Island . Speci men seen: Hertel 24 557. Gyalideopsis subantarctica Henssen and Lumbsch

part. Spores 8 in asci, uncoloured, simple, ellipsoid to curved, 9-1 1 x 3.5-6 ~m . Conidia rod -shaped , ca 3 x 1~m . TLC: one undetermined substance Ecology: on rock alongside streams, not far above wa ter

level. Distribution : Marion Island , Kerguelen .

Specimen seen : holotype (MB), Henssen 28 114d, 28 174a. Comments : a more detailed description can be found in

Henssen (1985). Lecanographa sp. Thallus extremely thin, crustose , effuse , pale ochre-grey, 1-2cm wide . Ascomata up to 0.5mm wide, either as single

apothecia , or most often apothecia merging together in a stroma-like structu re . No thalline excipulum. Excipulum

pro~

01/sledal and Gremmen

560

prium and hypothecium , including stroma-like tissue , brown-

crystals in apothecia, and presence of thallus areolae.

black, composed of a textura intricata. Hymenium colour-

less, 90-100~m high, uppermost part blue-black. Asci s[ender, ca 40 x 8~m, tip of the grumulosa type (see Egea and Torriente 1994). Spores 8 in asci , symmetrical , 27-32 x 2.5-3~m , mostly wi th 7 septa, uncoloured. Paraph yses slender, ramified to anastomosing. Pycnidia not seen. TLC: not performed due to extremely scanty thallus; thallus C+. Eco[ogy: on coastal plain , together with Carbonea spp. Specimen seen: Henssen 28 203, Marion [sl. Comments: it is not easily identified with any of the species treated in Egea and Torrente (1994) ; perhaps it comes closest to L. dime/aenoides (Egea and Torrente) Egea and Torrente from California and Mexico, but that species has a much better devetoped tha llus, and is also found in a much wa rmer climate.

Lecidea lapicida (Ach.) Ach. Thallus crustose , rimose to areolate, medium th ick , grey to buff-coloured, sometimes rustcoloured, up to ca 4cm diameter; prothallus black , thin. Apothecia black, sessile, up to 1.1mm diameter, margin distinct and persistent. Hymenium colourless, 50-70~m high, epithecium brown-green. Excip[e colourless in inner part. Hypothecium brown. Paraphyses branched to anastomosing , end ce[1 slightly enlarged, 3-4~m diameter. Spores el[ipsoid, 9-13.5 x 5-7~m . TLC: Stictic acid. Ecology: On rock outcrops in lowland mires; not ornithocoprophilous. Distribution:

Widespread

in

Northern

and

Southern

Hemispheres. South Georgia , Marion lsI. , South Orkney lsI. , Antarctic Peninsula . Specimens not seen , description based on Hertel (1984).

Lecanora disjungenda (Crombie) Herte[ and Rambold Thallus crustose, 3-4cm wide or more, white to grey,

smooth to rimose. Apothecia sunk in thallus, 0.5-0.7mm wide , excipulum proprium or thaHinum not evident, disc

black to brown-black, s[ightly concave. Hymenium 40--45~m high, epithecium brown-green . Spores broadly e[lipsoid, ca 12 x 7~m . Hypothecium pale yellow-brown . Excipulum proprium very thin , colourl ess. Paraphyses stout, simple , end cell not distinctly enlarged . Conidia threadlike, 10-18 x 1~m. TLC: alranorin and triterpenes. Eco[ogy: on black and grey lava rock, from low to medium elevations. Distribution: Kerguelen; Marion Island. Specimen seen: Gremmen 93-008 , 93- 009 , 93-010, 94-002, 94-62 , 94-071 , 94-072 , 94- 073. Lecanora flotowiana Spreng. Thallus as small (0.2mm) convex, green-grey areolae around the apothecia. Apothecia thick, with constricted base, up to 0.7mm diameter. Excipu[um thallinum thin, with crenulated margin. Disc fla t, warm chestnut brown . Excipulum thal[inum ca 80~m broad, with a very th in cortex. Excipu[um proprium 5- 10~m , colour[ess, composed of short, strongly agglutinated hyphae running parallel with paraphyses. Hymenium ca 40~m high, epithecium dislinct, brown. Spores 8 in asci, broadly elliptic, 9-11 x 6-7~m , with thick wa ll, Paraphyses stout, somewhat ramir,ed , end cell enlarged to ca 3~m , embedded in mUCilage. Very few small crystals in [ower part of subhymenia[ medulla . Pycnidia not seen. TLC: negative. Ecology: on lichens, bone algae and cyanobacteria, in supralittoral, together with Amandinea subplicata, Ca/oplaca sublobulata etc. Distribution : Marion lsI. , cosmopolitan. Specimens seen: Gremmen 93- 014, 94-263, Henssen 28068. Comments: the Marion Island specimens are fairly similar to the Western Antarctic populations of L. I/otowiana (0vstedal and Smith 2001), but differ in a number of details: more compact apothecia ([arger height/width ratio), thinner thalline margin, smaller spores and lower hymenium, fewer

Lecidea Iygomma Nyl. Thallus crustose, effuse, white-grey, rimose , medium thick. Apothecia up to 3mm diameter, immersed in thallus (rare[y ha[f-sessile), black, wi th a very thin excip[e which is surrounded by a carbonised margin of cortex origin. Hymenium 65-85~m high. Hypothecium brown. Spores 9-16 x 4-7~m. Medulla K/[ -. TLC: Norstictic acid. Ecology: On permanently damp grey lava rocks from [owland to high elevations. Distribution: Australia, South Georgia, Marion lSI. , Prince Edward Is[ ., lies Kergue[ en . Specimen seen: Gremmen 93-008, 94-193, 94 - 194, 94- 211 . Lecidea medusula (Dodge) Herte[ syn . Zosterodiscus russellii Hertel Thallus composite, up to 5cm wide, composed of numerous, dispersed, thin whitish areolae up to 1.5mm broad. Apothecia numerous, black, 0.5mm broad (rare[y up to O.7mm), half immersed in thal [us or rarely sessile with broad base, excipu[um proprium broad, split by a ring- like fu rrow into a massive outer part and a th in in ner part. Hymenium 45-70~m high, epithecium green-black. Hypothecium brown-black. Spores 8 in asci, simple , colourless , 9-14 x 4-7~m. Paraphyses simple or somewhat bran ched , end cells somewhat enlarged to 2.5~m diame ter. Pycnidia not Observed. TLC : negative. Ecology: on rocks from low to medium elevation. Distribution: South Georgia , lies Kerguelen, Marion Island , Prince Edwa rd Island . Specime ns seen: Gremmen 93-010, 94-71, 94- 73, 94-252. Lepraria neglecta (Nyl.) Lettau Thallus as 2-3mm wide patches, sometimes coalescing into larger patches , whi te to off-white, with a te ndency to margina[ lobes and medu[la. Soredia 109 ± 8~m diameter (95- 120~m), agglomerating into consoredia, which are 3-4 times as large. Soredium surface fluffy with hyphae ends.

South African Journal of Botany 2001. 67 552-572

561

TLC: atranorin , psoromic and roccellic (trace) acids Ecology: on Pannaria pulvinula . Distribu tion: Marion Island , cosmopol itan .

Notolecidea subcontinua (Nyl.) Hertel Thallus effuse , up to 3-4cm diameter, th in, appearing

Specimen seen : Gremmen 94-19.

to ferrug inous. Apothecia half sunk in Ihallus (Aspicilia-like), up to O.6mm diameter, margin black, pro truding , disc brown black, concave. Excipulum proprium ca. 100~m broad, dark

Leptogium tasmanicum F. Wilson Thallu s foliose, somewhat pulvinate, grey, sometimes tinted

smooth and continuous when wet , rimose when dry, whitish

brown in outer part, pale in inner part, composed of hyphae 80-90~m

brownish, 1-2cm broad. Lobes round to oblong, 2-3mm

wh ich originate in the medulla. Hymenium

wide , lobulate , lobules squamuliform . Apothecia not seen.

epithecium brown. Hypothecium pale brown. Asci 8-spored ,

TLC: not performed.

clavate, of Porpidia-type. Spores colourless, Simple, elliptic, halonate, 12-14 x 7-8~m. Paraphyses ram ified to slightly anastomosing. Pycnoconidia 4-8 x 1~m . Medulla KlI- . TLC: Atranorin .

Ecology: among mosses on rock at medium elevation. Distribution : Tasma nia, New Zealand, Prince Edward Island .

Specimens seen: Gremmen 94-236.

high,

Ecology: On wet rocks at low to medium elevation .

Lithographa marionensis Hertel and Rambold Thallus crustose , effuse , grey, thin,

rimose -areolate.

Apotehcia lirellate to 3-4-angeled, black, with raised excipu -

Dis tribution: An tarctic endemic: Sou th Georgia , lies Kerguel en, Marion lsI., James Ross lsI. Specimen seen: type (8M)

lum proprium and very narrow disc , O.8-1 .0mm long,

0.3-0.4mm broad. Excipulum proprium 100-1 30 ~ m broad, brown-black Ihroughou!. Hymenium ca. 1OO~m high. Asci broadly clavate, of Rimularia-type. Spores 8 in asci , simple, co lourless , ca . 12 x 6j.1m . TLC: gyrophoric acid.

up to 1mm broad , wax-like , pale orange. Apothecia sunk in inner areola te , opening with a small pore. Hymenium ca .

Ecology: on grey lava rock in feldmark at medium elevation. Distribution : Marion Island .

um colourless, composed of short, irregularly orien ted,

Literature: Hertel and Rambold 1990. Specimens seen : Gremmen 94- 193, 93- 003.

strongly conglutinated hyphae . Asci cylindrical. Spores uniseriate, uncoloured, simple, 26- 30 x 14-17~m , with double

Orceolina kerguelensis (Tuck.) Hertel Thall us placoid, orb icular, thick , in inner part strongly areo-

late, towards margin with elongate lobes, ca. 2mm long and

250~m

high, colourless. Excipulum proprium and hypotheci-

wal l. Paraphyses thin , f1exuose , anastomosi ng .

. Lithographa subantarctica Hertel and Rambold Thallus crustose, effuse, very thin , ochre to greyish , usually with rust-coloured part. Apothecia lirellate to 3-4-angeled, black, with ra ised proper margin and very narrow disc,

0.4-0.6mm long , 0.2- 0.3mm broad. TLC: negative . Ecology: on rock , from lowland to medium elevation Distrib ution: Marion Island.

Specimens seen : Gremmen 94-47 , 94-193, 94-21 t , 94- 214. Comments: this taxon is rather similar to L. marionensis, but differs in the usually rust-coloured thallus that lacks lichen products, the smaller apolhecia and the spores with halo. Massalongia carnosa (Dicks.) Korber Thallus foliose , up to 1cm diameter, usually developing isidia on margins and ra rely on lam ina, brown. Apothecia rare , laminal, sessile. Spores uncoloured, 3-septate, constricted

at septa, 16-26 x 6-7~m . TLC : negative. Ecology: on epilithic mosses (Andreaea sp.) from lowland to medium elevations. Distribution: Marion lsI., cosmopolitan in co lder areas .

TLC: one unidentified compound. Ecology: on rock in feldmark at medium to high elevations. Distribu tion: Kerguelen , Marion Island, Prince Edward Island . Specimen seen: Gremmen 93- 001, 93-006, 94-180. Pannaria dichroa Ny!. Thallus rosette-forming , to ca . 4cm wide , narrow and

cracked in inner part, flabellate and expanding to ca 3mm at margi n, pale grey-brown, slightly convex, up to 0.7mm high, longitudinally striated . Upper cortex paraplechtenchymatous, 50 - 60~m high , cells ± thinwalled, lumina ca 8 ~ m . Medulla dense, with strands of hyphae and clusters of Nostoc. Lower side weakly prosoplechte nchymatous in old parts with hyphae ends carbonised , appearing as a black rhizohyphae layer, 1 0- 20 ~ m high. Apothecia with thick thalline margin, when young almost globose with small concave disc and strongly res tricted below, when old up to ca.

3mm wide, with flat, brown disc. Spores 8 in asci, subglobose to broadly elliptic, 15-17 x 1 2- 1 4~m , with distinct epispor. Paraphyses simple, with constricted septa . TLC: negalive. Ecology: on rock.

Specimens seen : Gremmen 94- 001, 94-20.

Distribution : Kerguelen, Marion Island , Prince Edward

Comments: European and western An tarctic specimens

Island . Specimens seen: Hertel 24131 (M).

almost always have 1-septated spores, while the other known species in the genus, M. rnicrophylfiza (Ny!. ex Hasse) Henssen , growing on sandstone in California , has

2-3-septated spores (Henssen 1963). More material is necessary to clarify the situation .

Pannaria pulvinula P.M .J0rgensen T hallus up to ca. 10mm in diameter, pale och re , growing as rosettes , marginal lobes fiabellale, up to ca. 0.5mm broad, white-striated, with somewhat fluffy margin. In vertica l tran-

sect the thallus is

330-350~m

high. The upper cortex is hya-

0vsledaJ and Gremmen

562

line, Iwo-I ayered, uppermost layer ca. 1 5~m high, composed of a very compressed pseudoparenchymatous ti ssue, lumina slit-like, parallel to surface. Below that a ca. 100~m high pseudoparenchymatous tissue with rectangular, anticlinal lumina , and below thai a dense medulla , ca. 200~m high, composed of a very loose prosoplechtenchymatous tis sue. Hyphae colourless except the outermost ones which are more conglutinated and blue-black, with a few blue-black rhizi nes, ca . 3lJm in diameter and 25lJm long, with rough surface. Apothecia not seen. TLC : unidentified triterpenes. Ecology: growing on bryophytes (Andreaea sp.) at medium elenations; never collected on rock. Distribution: Marion and Prince Edward Islands, Kerguelen ,

Parmelia sulcata Taylor Thallus foliose, grey, 3-4cm wide. Lobes 0.7-1 .3mm wide, brownish at tips, with pseudocyphellae which develop into soralia on lamina. Lower surface black, with simple, black rhizinae. TLC: salazinic (major), consalazinic (minor), secalonic A (minor), atranorin (submajar) secalonic C (minor), strepsilin (minor) (94-122 J. Elix det). Ecology: on rock, growing over bryophytes, or epiphytic on other lichens, at medium elevation. Distri bution: Marion Island , Prince Edward Island , cosmopolitan. Specimens seen: Gremmen 94-122, 98-154, Henssen 28098.

Crozet

Specimens seen: Gremmen 94-18, 94 - 19, 94-240, 94-22. Comments: this characteristic species is not found with apothecia on Marion and Prince Edward Islands. Parmelia kerguelensis Wilson Thallus foliose , yellowish-grey or grey-brown, rosettes usually 3- 4cm diameter. Lobes 1.2-1 .8mm broad , with pseudocyphellae and isidia. Isidia barrel-shaped . Lower surface black, with simple rhizinae. TLC: atranorin, protocetraric acid. Ecology: on rock, growing over bryophytes, or epiphytic on other lichens, at medium elevations. Distribution : Marion Island , Prince Edward Island , Kerguelen . Specimens seen: Henssen ' 28063, Gremmen 94- 016, 94- 35,94-136,94- 012 ,98-154. Comments: this taxon is close to P saxatilis. Some authors have included it in that species. However, our material differs from P saxatilis from An tarctica and western subAntarctica in two characters: (1) the chemis try (P saxatifis has atranorin and salazinic acid) and (2) the form of the isidia (P saxatilis has cylindrical isidia; see Figure 5).

A

0.3 mm

B

Figure 5: Isidia of Parmelia kerguelensis (A) and P saxatills (8 )

Peltigera kerguelensis Dodge Thallus foliose, 5-6cm wide, lobs 10-12mm wide. concave . grey-brown . hairy. Lower side pale ochre , with a network of raised. pale veins. Apothecia common, on somewhat raised lobes, brown, slightly concave. TLC: negative. Ecology: on wet peat at entrance of bird burrow. Dis tribution: Kerguelen , Marion Island. Specimen seen: Gremmen 94-216 (0. Vitikainen deL) Peltigera truculenta De NoL Thallus foliose, 3-4cm wide, lobes 5-6mm wide, rounded , blue-grey, smooth. Lower side ~ uniformly pale, without distinct ve ins but with paler, uncorticated areas; rh izinae pale. Apothecia common, on stout ascending lobes, dark brown, saddle-shaped. TLC : gyrophoric acid , methyl gyrophorate, tenuiorin , pelti dactylin, dolicharrhizin , zearin, t5 (0. Vitikai nen det). Ecology: among mosses in sheltered sites. Distribution: Southern South America, Marion lsI. , Prince Edward lsI. Specimens seen: Gremmen 94- 11 4, 94-40, 94- 94, 94-65, 94-64 , 94-258,98-142 (0. Vitikainen deL).

Pertusaria cineraria Ny!. Thallus very thin, effuse, grey, scabrose, cracked. Apothecia sunk in thallus, 0.2-0.3mm wide, excipulum thallinum not visible , disc concave, ca 0.1 mm wide , black. Excipulum proprium 20-30~m wide , uncoloured. Hymen ium 90-100~m high, epilhecium brown-black. Spores 8 in asci, 30-34 x 17- 19~m , double-walled. Paraphyses thin, ftexuose, simple, strongly conglutinated , end cell not enlarged. Pycnidia not seen. TLC : negalive. Ecology: on rock in lowland feldmark . Distibution: Kerguelen. Prince Edward Island. Specimens seen: type (H-Nylander 23007) , Gremmen 94-135. Comments: the specimen has been compared to the type, and found to be similar in all essential details. The chemical content of the type, however, has not been investigated, as it is very small. The species has apparently not been collected since Nylander's time, and is not included in Archer's (1997) comprehensive list of Per/usaria species.

South African Journal of BOlany 2001 67 552- 572

563

Pertusaria werlhH Zahlbr. Tha llus off-white to pale grey, apothecia ve rr uciform, 0.8- 1.5mm in diameter, slightly constricted at the base. Os tioles black punctiform in a grey transluce nt zone, 1(-2)

Spores 8 in asci , simple , uncoloured , halonate , 18-30 x

per verruca . Ascospore s ellipsoid, smooth : 3-4 immature ascospores per ascus , becoming 3 mature ascospores per

8 -1 4~m . Paraphyses anastomosing . TlC: stictic acid.

ascus, (87-) 100-120~m long, 35-40 (-45)~m wide. TlC: 4-chlorolichexanthone (m ajor) , 4,5-dichlorolichexanthane (minor), methyl 2-0-methylperlatolate (minor), 2-0methylperlatolic acid (trace), 2-0-methylstenosporic acid (trace) together with traces of unknowns (J .A. Elix de!.). Ecology: on rock in saltspray zone.

Ecology: growing on small stones lying on wet soil in seepage areas.

Distribution: Marion lsI. , Kerguelen Specimen seen: Gremmen 94-42.

1.1 mm diameter, margin thick. Excipulum thallinum not present. Excipulum proprium thick , colourless in inner part.

Hypothecium colourless . Hyme nium

80- 130 ~m

high.

Distribution: Kerguelen, Prince Edward Island . Specimen not seen, description taken from Hertel (1984). Poeltidea perusta (Nyl.) Hertel and Hafellner Syn . Lecidea perusta Nyl. Thallus crustose, effuse , areolate, shiny brown. Apothecia

Comments: De!. R.S .Poulsen and U.S0chting .

black, up to 2mm diam ., half-immersed to sessile, with almost excluded margin. Hymenium 170-190 ~m. Excipulum

Pertusaria cf. perrimosa Ny!. Thallus crustose, effuse, thin, grey, cracked . Apothecial warts whitish , 1.2mm wide and 1mm high, conical. Ostiole partly black. Internal cavity ca 400~m wide. Spores 2 in asci, wi th double wa ll, 65- 75 x 35-4 5~m. TLC: stictic acid + unknowns.

Ecotogy: on rock in lowland feldmark.

proprium brown-black in outer part, colourless in inner part.

Hypothecium colourless . Paraphyses strongly agglutinated, anastomosing, end cell slightly enlarged . Spores 24-36 x 11 - 18~m , becoming greenish to brown when old , halonate. Conidia cylindrical, 4-6 x 1 - 1.5~m. Medulla KlI -. TlC: negative. Ecology: on rock. 3~-350m .

Distribution: Marion lsI. , New Zealand.

Distribution: Antarctic endemic:

Specimen seen: Gremmen 94- 135.

Kerguelen, Prince Edward Is ., South Orkney lsI. Specimen examined: Henssen 28131b .

Comments: only very few apothecial warts present , so the determination is not certain.

Phyllisciella marionensis Henssen Thallus umbilicate, dark red , outline often irregu lar, margin

effuse to effigurate , up to 10mm diameter. Photobiont cyanobacteria (Entophysalidaceae) with red-coloured gelatine sheath, in zones. Apothecia deeply immersed, urceolate , ostiote irregular, ca 0.1 mm across . Hymenium

South

Georgia , lies

Poeltinula cerebrinella (Nyl.) 0vstedal comb. nov. Basionym : Lecidea cerebrinella Nyl. , Nyl. in Crombie 1877 J. Un n. Soc. l ondon, J. Bo!. 15: 191. Thallus crustose, effuse, very thin , grey to green-grey, strongly cracked. Apothecia stromatoid, black, strongly contorted, constricted at base, black, up to 0.8mm diameter, convex. Discs lirelliform-irregular within the stromatoid

high. Paraphyses branched and anastomosing.

apothecium. Hymenium ca 1OO~m high, uppermost part yel-

Spores 8-32 in asci , simple , uncoloured. Conidia needle-

low-brow n. Paraphyses thin , anastomosing , end eel! not enlarged. Asci of Rhizocarpon-type . Spores 8 in asci , brown,

110- 170~m

shaped , 18-25 x 0.6~m . TlC: not performed. Ecology: overgrowing plant debris and bryophytes in upland feldmark. Distribution: Marion Island, Prince Edward Island Specimen seen: Gremmen 94- 214. Comments: only sterile specimens were found in Gremmen co llection. The description of apothecia is taken from

Henssen and BOdel (1984).

1-septated , constricted at septum, th inwalled 19.0 10.2 ~ 0 . 40~m (n=20). Pycnidia not seen. TlC: negative.

~

0.89 x

Ecology: on rock at medium elevation.

Distribution: Kerguelen, Marion Island. Specimens seen: Henssen 28148, Gremmen 94-73, type (H) . Comments: the apothecia may look similar to extreme forms

strongty rimose, with cephalodia. Cephalodia dome-shaped,

of Amandinea subplicata, but the latter species has a thicker and areolate thal lus, simple paraphyses and smaller spores, in addition to a different type of asci. The type (Nylander 10603, H) from Kerguelen consists of only two

2-3mm wide, pink. Apothecia aspiclloid, up to 2mm wide,

loose apothecia. No microscopic preparates were studied,

P/acopsis macrophtha/ma (Hook. and Tayl.) Nyl. Thallus subplacoid, thick , off-white , several cm diameter,

disc concave, flesh-coloured. Hymenium Spores 8 in asci, 18-20 x 10- 13~m. TlC: gyrophoric acid. Ecology: on rock in upland feldmark .

110- 120~m

high.

Distribution: Marion , Prince Edwa rd Island , Kergue len.

Specimens seen : Gremmen 94-239 , 94- 180, 93-029, 94-213, 94-182, 93-004, 94- 130. Poeltiaria urbanskyana (Zahlbr.) Hertel Thallus crustose, thin , white. Apothecia sessile, black, up to

but externally they appear similar to the collections from Marion Island. Porina leptalea (Durieu and Man!.) A.L. Sm . Thallus very thin , smooth, pate ochre, effuse , with TrentepohOa. Perithecia semiglobose , dark brown, ca 0.2mm diameter, in transect with a brown involucrellum in

upper part, excipulum uncoloured. Asci ca 55 x 1O~m, at tip thin, Kli -, without ocular chamber. Spores 3-septate, 20- 24 x 3- 3.5~m uncoloured. Paraphyses perSistent, simple, flex-

0 vsledal and Gremmen

564

uose, ca 1.5~m thick. TLC: negative. Ecology: on bird perching rocks and on rocks near the coast. Distribution: Marion Island , Prince Edward Island, Europe. Specimen seen : Gremmen 93-002, 94- 135, 94- 70. Pseudocyphel/aria crocata (L.) Vain. Thallus foliose , lobes broad , round . Upper side pale browngrey, ridged, with yellow saralia on ridges and lamina. Lower side pale brownish , tomentose, with scattered yellow pseudocyphellae. TLC: not performed . Ecology: among bryophytes in lowland , away from coast. Distribution: Europe, New Zealand, Prince Edward Island (also reported by Lindsay 1977). Specimen seen: Gremmen 98-137a. Psoroma asperellum Nyl.

Thallus squamuJose, squamules scattered, ascending, irregularely ramified, 2- 3mm long, 0.3-0Amm broad , upper surface pale green-brown, lower surface whitish to pale brownish, corticated. Prothallus not observed. Apothecia common, up to 1.3mm diameter, disc concave to flat , warm brown. Thalline margin "regular, lobulate. Cephalodia small , on margin of squamu les , blackish to dark brown. Hymen ium ca. 1OO~m high, epithecium brownish Spores 8 in asci , 18-25 x 12 - 14~m, epispore strongly warted. Pycnidia not seen. TLC : not performed. Ecology: among bryophytes on recent (1980) volcan ic ash deposit Distribution: New Zealand , Marion Island. Specimen seen: Gremmen 98-103. Ramonia subantarctica 0vstedal sp. nov. (Figure 6) Ramoniae nigrae similis, sed thalia saxicola et sporis latioribus. Holotypus: vide infra. Thallus crustose, very thin , grey brown , with Trentepohlia. Apothecia at firs t almost globose, with a small depression in the centre, later becoming urceolate with a small pore in centre, 0.3mm wide and as high, brown -black to black. Hymenium ca 100~m high . Excipulum proprium ca 90- 1101-/m wide , colourless excep t marginal areas , composed of angu lar, thinwalled cells, 6-8~m wide. Subhymenial part 100 - 150~m high , uppermosl part colourless, becoming brownish further down ; uppermost part composed of angular, th inwalled cells, further down becoming a textura intricata. Asci cyl indrical, 50 x lO~m , thickwalled , no Kli reaction observed. Paraphyses simple. Spores 8 in asci, uniseriate, 18-22 x 12- 15~m , uncoloured but when old fain blue-brown, muriform, not halonate . Periphysoids uncoloured, septated, 14-18~m long . TLC: not performed . Ecology: on rock , with Lecidea Iygomma, Lit/lOgrapha subantarctica and other lichens in upland feldma rk. Specimen seen: Gremmen no. 94- 252, Marion Island , Tafelberg , 240m a.s.l. , on Andreaea moss cushions in upland feldmark, 14.5.1994 (holotype BG). Comments: this species has most of the characteristics of the gen us, Le . urceolate apothecia, Trentepohlia as photobiont, angu lar, thinwalied cells in excipulum , simple para-

physes and the presence of periphyses (Coppins 1987, Thor and Vezda 1984, Vezda 1966, 1967, 1973). Th e growth on rock is unusual , but one other species (from California, see Coppins 1987) has also been found on th is substrate. It is somewhat similar to R. nigra from Great Brita in, but the spores are relative ly much broader and the ecology different. RIJizocarpon geographicum (L.) DC. Thallus crustose, effuse, up to 7cm diameter, several often coalescing to a continuous colony up to 20cm across, yellow-green, areolate ; areolate round , convex, up to O.8mm wide, on a black hypothallus, prothallus black, narrow. Apothecia black, round to angular, up to 1mm diameter, nat with a slightly protruding proper margin when young, convex with excluded margin when old. Hymenium 100-120~m high , epithecium brown with a violet tinge, K+ red . Spores 8 in asci, early dark, 30- 34 x 12-14~m, with 3 longitudinal septa and 3--4 transverse septa. Medulla KlI + blue . TLC : Rh izocarpic, ~ psoromic acids. Ecology: on rock from low to medium elevation. Distribution: Marion lsI. , cosmopolitan. Speci mens seen: Gremmen 94- 60, 94-73 , 94- 88 Rhizocarpon nidificum (Hue) Darb. Thallus crustose, effuse, up to 2cm diameter, usually between 1 and 1.5cm , composed of pale green areolae dispersed on a thick, black prothallus. Areolae flal to somewhat

E E

ci

B

c

E

"-

o

N

Figure 6: Ramonia subantarctica. A: vertical secian of apothecium: S' ascus wi th paraphyses. Ho!otype

South African Journal of Botany 2001 . 67' 552- 572

convex , O.1-0 .2mm diameter, distance between areolae ca . 3 times diameter of areolae. Apothecia black , angula te due to compression, 0.3-0.5 (-O.7)mm diameter, margin raised also when old, shiny; disc flat, rough , matt. Hymenium 60 - 70~m high, colourless to pale viole I, K+ burgundy red, epithecium brown-red , K+ burgundy red , no crystals seen. Spores 8 in asci , early dark greenish brown, mostly 3-seplate (ca. 80% 3-septate, ca. 20% 1-2 -sep tated , ca. 30% of all wilh 1 longitudinal septum), 15-30 x 8-13~m. Medulla K/I+ blue. TLC: Rhizocarpic acid. Ecology: on rock at medium elevation. Distribution: Antarclic Peninsula , South Shetland lsI. , South Georgia , Marion Island. Specimen seen: Gremmen 94-72.

Rhizocarpon poslumum (Nyl.) Arnold Thallus 4-5mm wide, in mosaics with other lichens , with 0.3- 0.4mm broad grey-brown areolae imbedded in a black prothallus. Apothecia numerous, black, slightly elevated above thallus, thin, flat , up to 0.5mm diameter, w ith excipu lum proprium slightly elevated. Hymenium 170-180~m high , epithecium prominent , wi th brown-red granules and also a green-black pigment. Excipulum proprium 80-90~m broad, red-brown in outer part. Hypothecium dark brown. Asci with 8 spores. Spores persistently colourless, 18-22 x 8- 10~m , most with three transverse septa, but also a few submuriform or with only two septa . Paraphyses anastomosing . Pycnidia not seen . TLC : negative . Ecology: on rocks at medium eleva tion. Distribution: N Europe, Canada (?) (Purvis el al. 1992), Marion Island . Specimen seen : Gremmen 94-73, 94-66 . (aff.) Rinodina griseosorafifera Coppins Thallus evanescent. emerging as soralia , which are up to O.5mm in diameter, convex , coa rse, pale grey-green. Soredia compact, 20-25~m diameter. K -. No apothecia or pycnidia seen. TLC : atranorin, zeerin . Ecology: on exposed stems of Azoreffa; med ium to high elevations. Distribution: Europe , N America , Marion Island, Prince Edward Island. Specimens seen: Gremmen 98-103, 94-236. Comments: According to T0nsberg (pers. comm.) the specimen shows some similarities wi th the chem ically concordant R. griseosoralifera , a species fairly common in NW Europe, but never before recorded from the southern hemisphere.

Rinodina peloleuca (Nyl.) MOil. Arg. Thallus crustose, thick, pale grey to brownish , rimose. Apothecia sessile, up to 0.5mm wide, with thalline margin, concolorous with thallus , disc red- brown. Hymenium 100-1 1 5~m high, hypothecium uncoloured. Spores 8 in asci, of Physconia-lype, 15-23 x 10-13~m . Paraphyses simple, end cell up to 6 ~m diameter. Conidia bacilliform , 4 x 1.5~m . TLC: skyrin, triterpenes .

565

Ecology: on coastal rocks. Dis tribution: Prince Edward lsI. , circumaustra l (Matzer et a/. 1998). Specimens seen : Henssen 28096b, 28096c (H. Mayhofer det).

Scoliciosporum umbrinum (Ach .) Arnold Thallus crustose, scurfy-granular, cracked , brown -grey, partly rusty. Apothecia 0.2-0.3mm broad , sessi le, without or with very thin true margin, dark brown. Algae Trenlepohlia? Hymenium pale blue. 35-40~m high , epithecium strongly blue-green. Excipulum proprium blue-green , composed of radiating hyphae. Paraphyses ramified , end cell not enlarged. Spores 7- 9-septated , uncoloured, 17-20 x 2~m , twisted in asci. TLC: not performed. Ecology: on rock. Distribution: Marion lsI. , Europe, Australia . Specimen seen: Henssen 28 114 c. Comments: S. compactum Korber from Australia. differs from typica l S. umbrinum in having a blue-green epithecium, but according to Purvis el al. (1992) th is character is not reliable. Thus we have kep t the name S. umbrinum. Siphulaslrum mamillalum (Hook and Taylor) D. Galloway Thallus squamulose, up to ca. 1cm in diameter, squamules covered by terete to somewhat flattened , simple to coralloid. greyish black papillae; at margin squamules nattened , enlarged, brownish . Apolhecia up to 3mm wide, thin sessile, with f1exuose outline, thalline margin absent , proper margin thin, protruding , somewhat paler than disc; disc dark brown. Hymenium 110 -120~m high , brown in uppermost part. Hypothecium pale brown. Excipulum proprium composed of a subpseudoparenchymatous tissue, with cell lumina elongated and radia ti ng , hyaline except outermos( ones which are brownish. Spores 8 in asci , ellipsoid with pointed ends and rough surface, ca 16 x 5~m. Paraphyses little ramified, thin, fl exuose. end cell slightly enla rged to 2-2.5 ~m . TLC: pannarin . Ecology: epiphytic on epilithic moss cushions (Andreaea sp., Racomitrium sp.) at medium elevations. Distribution: Marion lsI. , Prince Edward lsI., circumaustral. Specimens seen: Marion Island: Gremmen 94-21 , 94- 177, 94-235. Sporaslalia tesludinea (Ach.) A.Massal. Thallus subeffigurate, growing in rosettes ca. 1cm diameter, brown , shiny, areolate , areolae at marg in elongated. Prothallus narrow, black. Apothecia in inner part of thallus, black, between areolae, angular due to compression, up to O.4mm diameter, within thin , raised, proper margin and concave disc. Hymenium colourless, 130-140~m high. Asci clavate 45- 50 x 16 - 18 ~ m , polysporous. Spores globose, 2.5-3.5~m diameter. TLC: gyrophoric acid. Ecology: On rocks at medium elevation . Distribution : Central and Northern Europe, N. America , Asia , South Georgia , Antarctic Peninsula, Marion lsI. Specimen seen: Gremmen 94-214. Steinera sorediata P. James and Henssen

0vstedal and Gremmen

566

Thallus placoid, effigurate, grey to yellowish-grey, up to 2cm diameter, forming lobes .±. expanded at margins, inner part of thallus areolate. Saralia on lobe margins. Lower side ecorticate , blackish . Apothecia or pycnidia not seen on Prince Edward Islands material. TLC: negative. Ecology: on eroding lava rock. from low to high elevations. Distribution: New Zealand , Marion Island , Prince Edward Island. Specimens seen: Gremmen 94-243, 94-71 , 94-214 , 94-244. Comments: four species are at present known in this entirely austral genus (Henssen and James 1982). occurring from New Zealand to Kerguele n, and now for the first time record ed from th e Prince Edward Islands. Our specimens differ from the description of S. sorediata in that the soralia are marginal, as opposed to laminal in the description, and it may be an undescribed species, but this can only be ascertained on ferIile material. Stephanocyclos henssenianus Hertel Thallus very th in, cruslose, grey. Apolhecia black, up 10 3.5mm diameter, strongly gyrose. Excipulum thall inum not presen t, excipulum proprium and hypot hecrum black. Hymenium 70-105~m high. Asci of Porpidia-type. Spores 8 in asci, simple, uncoloured , halonate, 13-30 x 5.5-9~m. TLC : confluenl ic and 2'-0-methylperlatolic acids. Ecology: on rocks and loose stones at low to high elevation ; very common Distri bution: only found on Marion and Prince Edward Island . Specimens seen: Gremmen 93-005, 93- 051, 94-130.

Stereocaulon implexum Th.Fr. Thallus fruticose , lower part ca 1mm wide, uncorticated, pink , scabrous, upper part ramifi ed, corticated , green, grey with squamulose phyllocladia. Cephalodia numerous in

upper part, up to 0.6mm diameler, semiglobose, grey, wilh darker ridges so as to appear scrobiculate in vert ical transection with paraplechtenchymatous tissue between the Nostoc colonies. P.pothecia not seen. TLC: atranorin and stictic acid complex.

Ecology: between bryophyles on mossbank at high elevation. Distribution: Marion lsI., south ernmost South Amer ica,

Tristan da Cunha and Gough 151. (Lamb 1977, 1978). Specimen seen: Gremmen 98-86. Comments: specimens of this species with the same chem-

ical content have been found recently on Gough Island (Gremmen leg. ). TeplJrome/a atra (Huds.) Hafellner Thallus crus tose, effuse , thick, white , up to several cm diam-

eter. Apothecia with Ihick thalline margin which is concolorous with thallus, disc, flat, black. Hymenium ca 100~m high, faint violet , epithecium red-brown . Spores 8 in asci. simple, uncoloured, ca 12 x 6~m. Paraphyse end cell ca 5~m diameter, lumen 1-2~m . Hypothecium brown. TLC: atranorin, alpha-collatolic, alectoron ic acids. Ecology: on inland rocks , allow to medium elevation. Distribution : Marion Island, Prince Edward Island . cosmo -

politan. Specimens seen: Gremmen 94-70, 94- 73, 94-137 . Comments: a uniform species in the whole antarctic and sub-antarctic area .

Sticta fuliginosa (Hoffm .) Ach. Thallus foliose , with broad lobes, upper side brown , with isidia in cora lloid clusters on margin and lam ina. Lower surface pale brown, lomenlose, with scattered, pale cyphellae. TLC: negative. Ecology: among bryophytes in fern brake , with Pseudocyphellaria crocala. Distribution: Prince Edward lsi. , cosmopolitan , including

New Zealand (Galloway 1997). Specimen seen: Gremmen 98-137b. Siereocaulon cymosum Cromb. Pseudopodetia 2-3cm high , with a distinct main stem in lower part , but much divided in upper part. Main stem noncorticated, whitish, sometimes pale orange in lower part , smooth . Phyllociadia papil lose to verrucose, sometimes coalescent to crenate-squamulose. Cephalodia blue-grey,

TeplJromela eatonii (Cromb.) Hertel Thallus crustose, effuse. 2-3cm or more in diameter, grey-

white, as roundish areoles embedded in black prothallus, or more cohere nt and then rimose. Apothecia black , su nk in

thallus , without thalline margin , 0.3- 0.5mm diameter. Hymenium 45-55~m , epithecium blue-black. Hypothecium brown . Spores 8 in asci , simple, uncoloured , 9-11 x 4-6j..1m. Paraphyse end cell ca 4~m bro ad, lumen ca 1 ~ m broad. Pycnidia not seen.

TLC : Atranorin. Ecology: on inland rocks fro m low to high elevations, often together with Lecidea Iygomma. Specimens seen: Gremmen 93-010, 94-211, 94- 214, Henssen 28 041 b. Distribution : Marion Island, Kerguelen . Comments: our specimens have been compared to the type

cauliflower-like, shortly stipitate al base, sessile higher up on

from Kerguelen (8M) and were fou nd to be similar in all

pseudopodetia . Apothecia common , terminal , up to ca 2mm in diameter, margin strongly incurved and sculptured, disc

essential details.

concave , brown . Spores 3- 6-septate, 33-42 x 4 -5~m .

Thelenella kerguelena (Nyl.) H.Mayrhofer Syn. Microglaena austrocinerascens D.C. lindsay Thallus crustose , effuse , th in , pale grey to pale brown , often rimose. Perithecia scattered, up to O.5mm diameter, semi-

TLC: atranori n, lobaric acid with accessories. Ecology: on lava rocks and scoriae deposits, at low to medium elevations. Apparently a coloniser on lava outcrops. Distribution: Kerguelen, Marion Island , Prince Edward

Island . Specimens seen: Gremmen 93-005 , 94- 132, 94-88, 98-61 , 94- 133, 94-172,98-113.

globose, black, shiny. Involucrellum black, only present in upper part of perithecium. Spores colou rless , ellipsoid -elongate, 8 in asci, muriform, with 5-7 transverse and 3 longitu-

dinal septa, 20-32 x

10-16 ~m .

567

South African Journal of Botany 2001 . 67 552- 572

TLC : not investigated . Ecology: growing on inland rocks.

Distribution : Marion lsI., Kerguele n, Heard lsI., South Shetland lsI. Specimen not seen: description based on Antarctic speci ~ men.

Thelocarpon subantarcticum 0vstedal sp. nov. (Figure 7) T epiboli similis, sed pru ina destituta et sporis minoribus .

Holotypus: vide infra . Thallus apparently absent, apothecia embedded in a gelatinous

a lgal

crust.

Apothecia

at

first

pe r i the cia~like,

0.2-0.3mm across, globose , whitish , with semi-hyaline cen ter later on open ing up, c up ~ l i ke, to O.Smm diameter, disc concave, flesh ~ coloured . Hymenium uncoloured, ca 100IJm

high. Hymenial gelatine Kli -. Paraphyses , simple, strongly agglutinated . Excipulum proprium uncoloured , 30-40~m thick, composed of long , periclinal hyphae. Asci slightly bottle-shaped , 50-60 x 35-40~m, multi-spored (>100) , Kli -. Spores uncoloured, without pseudosepta, 1.5-2 x 1~m.

coloured; prothallus often conspicuous, black , Apothecia subimmersed to immersed , black , up to O.Smm diameter, margin well developed and pe rSistent. Hym en ium 90 - 12 5 ~m

high, upper part co lourless to green-grey.

Excipulum proprium

brow n ~ black .

Hypothecium dark brown .

Spores ellipsoid, 10-14 x 6-9~m. Pycnoconidia cylindrical, ca. 4 x 1~m . Medulla KlI-. TLC: Negative. Ecology: on rock Distribu tion: Cosmopolitan in cold reg ions. Antarctic: Sou th

Georgia, Marion lsI. (listed by Lindsay 1977 as Lecidea atrata (Ach.)Wahlenb.) , lsI. Kerguelen , South Orkney Is., South Shetland Is., Antarct ic Peninsula. Specimens seen: Gremmen 93-009, 93-0 10. Turgidosculum complicatulum (Nyl.) J. Kohlmeyer and E. Kohlmeyer Thallus 5-10mm high , brown -green . Perithecia 0.3mm in outer diameter, inner diameter ca. 170lJm . Most spores

TLC: not performed.

unseptated , but a few 1-septated 'spores occur. Spore size: 12.7 ± 0.47 x 4.2 ± 0.40~m . TLC : negative.

Ecology: on brittle tuff, with Steinera sorediata .

Ecology: in supralittoral on rocky shores, with Verrucaria

Specimen seen: N.J .M. Gremmen no 94-253, Prince Edward Island, eastern rim of kent Crater, 290m a.s.l., on brittle tuffaceous rock , 15.5.1994 (holotype, BG). Comments: in the British flora (Purvis et al. 1992), it keys out

other lichens. Distribution: Marion Island, Prince Edward lsI.,

Periphyses not seen.

durietzii, V. tesselatula , V. psychrophila, V. ceutocarpa and circumaus~

close to T epibolum. However, that species has yellow pru ~ ina , a hyme nium gel which is Kli + blue and larger spores.

tral, Arctic (?). Specimens seen : Gremmen 93-01 4, 94--271 , 93-050, 93- 016.

Tremolecia atrata (Ach.) Hertel

Comments: often called Mastodia tesse/ata .

T hallus crustose , effuse , th in, up to 2cm diameter, rust Usnea antarctica Du Rietz

Thallus fruticose, 2-5cm high yellow-green, usually from a delimited holdfast, erect, branched in upper part with numer-

O,5mm

A

ous attenuated branches . Branches terete , usually va riegat-

ed with black pigment. Surface matt, subpapillate to grossly papillate. Soralia usually common throughout thallus, plane to excavate , with cra teriform margin. Medulla compact; axis

thick. Apothecia very rare , subterminal, up to 10mm diamete r, concave and irregularly saddle~shaped , externa l surface wrinkled , wrinkles sorediate, no fibrils. Thalline margin thin,

irregular. Disc smooth to slightly wrinkled, pale brown. Spores simple, colourless, 8 in asci, ellipsoid , 9- 10 x 6-7~m .

TLC : usnic, fumarprotocetraric acid com plex. Ecology: on rock away from the coast, at low elevations.

c

B E

::l

o~I

o

'"

Distribution: New Zealand (South Isl.), southernmost South America. Antarctic: Prince Edward Island, South Georgia , Bouvet. ya, South Sandwich Is. , South Orkney Is., South Shetland Is., Antarctic Pe ninsula , continental Antarctica .

Specimens seen : Gremmen 98-155, 94- 135, Henssen 28099 . Usnea maculata Stirt.

Thallus fruticose, 2-8cm high, with a centra l hold fast, main bran ch up to 3mm broad , with numerous fibrillae and a few,

scatlered papillae. In term inal bran ches the papillae are numerous but breaking off quickly to leave scars looking like

pseudocyphellae. In one population (Gremmen 98- 148) Figure 7 : The/ocarpon subantarcticum. A: vertical transect of

apothecium: B: ascus with paraphyses: C: spore . Holotype

clusters of isidia-like pro tuberances emerge from these

568

scars. Al l branches red-maculated . In transect , the axis consti tutes 70- 80% of the main branch , and the re is a narrow.

dense medulla wi th algae. TLC: 1) with usnic and protocetraric acids (98-148); 2) usnic and stictic acid complex (94-15 and 98- 156). Ecology: on rock away from the coast, from low to medium elevations.

0vsledal and Gremmen

thalli. Perithecia emergent, smooth, black, up to O.2mm diameter. Involucrellum in upper part only. Excipulum brown

in upper part, pale in lower. Spores 8 in asci , ellipsoid. 7-12 x 4-5~m. Eco logy: on intertidal rocks, just above splash zo ne. Dis tribution : Marion lsI. , cosmopolita n in col d areas. Specimen seen: Gremmen 94- 272,

Distribution : Africa, New Zealand , Marion Island, Prince

Edward Island. Specimens seen : Gremmen 94-15, 98-148, 98-156 (P. Clerc de!.) . Comments: in Swinscow and Krog (1988) , Galloway (1985) and Stevens (1999) only the protocretraric acid chemotype is mentioned. Thus the slictic acid type is new.

Verrucaria ceutocarpa Wa hlenb. Thallus effuse, th in , black to brown-black, strongly areolate, a reoles 0.2-O.3mm across , fissures concolorous with thallus. Perithecia almost sunken in areoles , only black ostiole

protruding. Involucrellum only in upper part, excipulum co lourless. Spores 8 in asci, 9-12 x 6-7~m. Ecology: in tertidal rocks and rocks within the salt-spray zone. Distribution : Marion lsI., cosmopolitan in cold areas. Specimens seen: Gre mmen 94-270, 93-013 . Comments: no differences could be found between these specimens and those from the western Antarctic area

(0vstedal and Smith 200 1) or from N Europe (material in BG) .

Verrucaria ct. dltmarsica Erichsen Thallus very thin, 2-3mm wide, without prothallus, gelatinous, green-grey, with numerous black dots, ca. 0.02mm diameter, wh ich sometimes coalesce to uneven ridge like structures . Peri the cia half-sessile, 0.2-0.3mm in diameter, with rugulose su rface. Involucrellum entire ; excipulum pale .

Spores 8 in asci, regula r, ellipsoid, 8-9 x 5-6~m . Ecology: on intertidal rocks , togethe r with V lesse/alula , V, mawsonii and V. halizoa . Distribution : Marion lsI. , Europe. Specimen seen: Marion Island : Gremmen 94-272. Comments: Keys ou t as V ditmarsica in N European floras , but differs in the rugulose surface of the perithecia and the regu larily ellipsoid spores (kid ney- to D-shaped in NW European ones , see Purvis el a/. 1992) . Verruca ria durietzit' Lamb

Verruca ria mawsonii Dodge Thallus up to 1cm diameter, 0.2-0.3mm th ick, green-brown , strongly rimose . Rimae black, partly with raised edges. Areolae with small black dots (0.02-0.05mm) which some times coalesce to irregular ridges . Perithecia completely sunken in thalli ; inner diameter ca. 130jJm. Involucrellum entire, Irreg ular a nd expanded in upper part , thin and regular in lowe r pa rt . Excipulum colourless. Spores 8 in asci,

12-14 x 6 - 7~m. Ecology: just above splash zone, with Vumbilicala and Vpsychrophila , 7m above sea. Distribution: Marion Island, Heard lsI. Specimens seen: Gremmen 94-272 , 94- 265 , holotype (FHDodge). Comments: the specimens from Marion Island are fairly well in accorda nce w ith the type specimen from Heard lsI. ,

except that the type is less rimose at the marg in of the thallus and the areolae have only dots which do not coa lesce to ridges . However, in part of the thalli of the Marion Island specimens, the areolae have only dots, so this character is

apparently modified by exposure. Another possibility is Vdegelii R. Sant. , occurring in N Europe but that species has beUer developed ridges (Figure 8). Perhaps V mawsonii and V degelii belong to a variable, widely dispersed taxon, but until more specimens are studied, th is will only be speculation.

Verruca ria maura Wa hlenb. Thallus effuse, dark brown to black , medium thick, dotted, ri mose, cracks concolorous wi th thallus . Perithecia medium to much prot ruding , diameter 0.6-0.9mm , inner diameter 200- 300~m . Involucrellum entire, black, excipulum pale. Spores 8 in asci, ellipsoid , 16-19 x 8-13~m. Eco logy: in intertidal and supraliltoral rockpools, also inland, up to 250m above sea level.

Distribution : Marion ls I. , Prince Edward lsI. , cosmopolitan in cold areas.

Specimens seen : Gremmen 94- 267, 94-274 , 94-241 , 94-238.

T hallus effigura te , up to several cm in diameter, margins

thick, with radiating lobes separa ted by bla ck cracks, smooth, whitish. Perithecia immersed, up to 0 .3mm in diameter. Involucrell um entire, black; excipu lum pale. Spores 8 in

asci, 13- 15 x 8- 9~m . Ecology: rocks from itertidal to more inland , but withi n salt-

Verrucaria mucosa Wa hlenb. T hall us effuse . continuous, relatively thick , olive-green, smooth . Perithecia completely imme rsed , up to 0 .2mm diameter. Involucrellum only in upper pa rt, exci pulum uncoloured. Spores 8 in asci , ellipsoid, 9-13 x

6-7 ~ m .

spray zone . Distribution : Marion lsi., circumaustral .

Ecology: on rocks in intertidal rock poo l. Distribution: Prince Edward lsI., cosmopo litan in cold areas.

Specimens seen: Gremmen 93-014, 93- 013 , 93-016, 94-41 .

Specimen seen: Prince Edward Island: Gremmen 94-238.

Verrucaria ha/izoa Leight. Thallus effuse , thin , brown-green , smooth , usually small

Verruca ria psycrophi/a Lamb Thallus effuse, up to several em wide, brownish, of med ium thickness, cracked , with cracks irregu lar, not blackened.

South African Journal or Botany 2001 67, 552- 572

569

gins brown-black, :': shiny, 0.2-0Amm thick. Photobiont green algae , sim il ar to those found in V. e/aeochroma and V. tesse/atu/a. Perithecia black, semi-immersed , O.1-0.2mm

A

wide ,

semiglobose

with

small ,

impressed

ostiole,

Involucrell um black, reaching halfway down the perithecium.

Excipulum propri um dark brown in uppermost part, lower down uncoloured. Paraphyses soon disintegrating : periphy-

ses present, lax, ca, 15 x 1~m, Spores 8 in asci , broadly elliptic, 9- 10 x 5- 7]Jm. Pycnidia common on some thalli, almost

superficial, dark brown, 0,05-0,1 mm diameter. Conidia bacilliform, ca 2 x 0 . 5~m, uncoloured .

1 mm

Ecology: on grey lava rocks in or just above the intertidal

1 mm

zone, together with Vtesse/ata and V psychrophi/a, Distribution: on ly known from Marion Island.

Specimens seen: Gremmen , 94-265 , Marion Island, Macaroni Bay, 2m a.s.l., grey lava coasta l rocks, 7m from

shore, just above splash zone, 16.5.1994 (holotypus, 8G), Gremmen 93- 011, 93- 012, 94-264, Xanthoparme/ia mougeotii (Schaerer ex D. Dietr.) Hale

Thallus fotiose, appressed to substrate, orbicular, ca 1cm diameter, yellow-white, sorediate. Lobes spreading in an

intricate pattern, 0,2-0,3mm wide, up to 0.5mrn long. Soratia Figure 8: Thallus habitus of A: Verrucaria degeJii R.Sant. (Santesson 18849, Finnmark, Norway, BG); B: V mawsonii Dodge (hololype, BANZARE, B 140-2 , Heard Island, 1929, FH-Dodge): and C: V mawsonii (Marion Island, Macaroni Bay, Gremmen 94265, BG, GREMMEN)

Perithecia little to moderately protruding, visible part dark

lam inal , up to 0.6mm wide. Upper cortex brownish , ca 20].Jm

high, composed of a textura intricata, Medulla loose, wi th algae in upper part . Lower cortex pseudoparenchymatous

with thick-walled cells, brown,

10-15~m

high , with a few

simple, black rhizinae . Apothecia not seen. TLC : usnic, isousnic, stictic, and norstictic acids. Ecology: on rock, also overgrowing moribund mosses, medium elevation.

broad , brown-black . ExcipuJum colourless. Spores 8 in asci,

Distribution: Europe, Marion Island (also reported by Lindsay 1977), Specimen seen: Gremmen 94-067.

broadly ovate, 12-15 x 7-9~m . Ecology: Intertidal rocks , together with V tesse/atu/a, V

Xanthoria candelaria (L.) Th, Fr.

brown, ca. 0.1 mm in diameter. Inner diameter of perithecia

ca.

250~m.

Involucrellum on ly in upper part,

30-40~m

duritzii, V. umbilicata, etc.

Distribution : South Georgia , South Orkney lsI., South Shetland IsL, Antarctic Peninsula, Marion Island.

Specimens seen : Gremmen 93-027, 93-011 , 93-012, 94-264b, 93- 013, 94-4 1.

Thallus frut icose, ascending, lobes flat, much ramified at apex , orange, sorediate. Soralia on lower side of lobe ends . Soredia ca . 20].Jm diameter. Apothecia subterminal, not rare , up to 0.9mm diameter, basis thin , constricted, margin th in, disc flat, smooth , somewhat darker than thallus . Hymenium

diameter. Involucrellum on ly in upper part, excipulum co lour-

high . Spores 8 in asci, ca 12 x 6~m, septum Paraphyse end cell ca 3~m diameter. TLC: not performed, Ecology: on stems of Acaena magelfanica in lowland herbfield •

less , Spores 8 in asci, 12- 15 x 7- 9~m , Ecology: on rocks in the intertidal zone, and along the edges

Distribution: cosmopolitan. Specimen seen: Marion Island: Gremmen 93-042.

ca,

Verruca ria

tesselatula Ny!.

Thallus effuse, thick, grey-brown, areolate, cracks black, Perithecia emergent to ca 1/3, dark brown, up to 0.3mm

of coastal rock pools . Distribution : Marion lsI. , circumaustral.

Specimens seen: Gremmen 94- 272, 94- 270, 93-027 , 93-0 12, 94-264b, 93-013, 93-016 ,

50~m

1 - 3~m,

Zahlbrucknerella marionensis Henssen

Thallus filamentous, dark olivaceous to blackish, tufted , Filaments 3-5mm long ,up to 60~m thick, at base ca. 1OO ~ m thick. Hyphae comprising a basal strand of 1-3 rows of

Verrucaria umbilicata 0vstedal sp. nov. (Figure 9)

angular cells, in upper part there is a network of angular cells

Thallus subfoliosus, umbilicatus, ochraceo ~ griseus , saxicola. Perithecia semi-immersa, 0 .1-0.2mm lato, sporae 9-10 x

which are connected with thin connections. Photobiont

5-7~m ,

Holotypus: vide infra. Thallus 1-3mm wide, subfotiose, fastened to the rock in the

Ecology: adnate to rock, above water level in river.

centre and with margins free. Upper surface ochre-greyish, non- rimose, lower surface black, irregularly pappilose: mar-

Specimen not seen, description based on Henssen (1985)

Scytonema, trichomes twisted. Apothecia not observed .

Distribution: Marion Island.

Qlvstedal and Gremmen

570

bose , halfway covered by thalline tissue, black area ca. O.3mm in diameter. Involucrelium extensive , spreading laterally, excipulum black, entire . Spores 8 in asci, narrowly

ellipsoid , ca 20 x

6 ~m.

Periphyses ca. 10 x

0 . 5~m .

Ecology: on rock, together with Verrucaria mawsonii and V

durietzii. Distribution: Heard Island. Specimen seen : Holotype (FH-Dodge), Heard Island. Comments: The algae are similar to those of V. durietzii and other Antarctic Verrucaria species with thick tha lli, and are not cyanobacteria, as stated by Dodge (1948). Thus this species must be transferred to Verrucaria . It appears to be related to V durietzii, wh ich also has a very thick, black hypothallus, a thick, whi tish thallus, and an involucrellum which spreads laterally. However, V durietzii differs by a thallus that is placoid , and has longitudinal black cracks, Figure 9: Verrucaria umbilicata . Scale = 1mm . HoJotype

perithecia which are smaller and less protruding, smaller, relatively broader spores, and lacking pits on the surface. The nature of the pits , however, is not clear, maybe it is an

During the study of Verrucaria taxa , we came across the following two species , placed in other genera but belonging in Verrucaria , and we recombine them into this genus.

artefact ('Schneckenfra(>').

Verrucaria nigra (MOIl. Arg.) 0vstedal comb . nov.. Basionym: Dermatocarpon nigrum MOIl. Arg; MOIl. Arg. Mission Sci. Cape Horn 1888: 170. Thallus dimorphic; vegetative thallus sterile, dark brown , rimose, cracks black , individual areolae O.3- 0.4mm across ; in vertical section ca 100IJm high, composed by a pseudoparen chymatous tissu e with scattered photo bionts.

Secondary thallus at first as circular, slightly raised , black parts of the primary thallus, later enlarged up to 2.5mm, raised and free at the margins. The secondary thallus is fer-

tile, raised 0.1-0.2mm above the vegetative one, and attached to the surface by strands of carbonaceous tissue , which is interpreted as remains of a more continuous carbonaceous tissue . In vertical section ca . 250~m high, with prom inent carbonaceous tissue , which is weake r developed

in the central part and which has ridges towards upper and lower surfaces. Ridges towards upper surface ca . 30~m high and 8-15~m wide. Non-carbonaceous parts between

ridges filled with photobionts. Ridges seen on surface as a weakly ra ised, reticulate pattern. Photobionts ch lorococcoid (?) , ca 3~m diameter, often angular due to close packing . Perithecia on upper surface, flush with surface, centre

broadly obpyriform ,

15 0- 170~m .

No evident involucrellum,

but rays of carbonaceous tissue around the perithecium. Excipu lum ca. 20~m broad , colourless. No hamathecium

seen. Periphyses distinct. Asci subglobose , K11 -. Hymenial gelatine Kl i blue. spores 8 in asci , simple , colou rl ess, 6-7 x 4- 5~rn.

Distribution: Cape Horn.

Specimen seen: holotype (G). Verrucaria tesse/ata (Dodge) 0vstedal comb. nov. Basionym : Phyllopyrenia tesse/ata Dodge; Dodge BANZARE VII, 1948: 53 Thallus milky white, fairly th ick (up to ca 0.8mm), effuse, secondarily cracked , thal1ii ca 1cm diameter, margin not effigu rate, surface with numerous minute pits . Photobiont

green algae. Hypothallus thick, black. Perilhecia semiglo-

Discussion The present account lists 100 species of lichens for Marion

and Prince Edward Islands, and includes, besides the species that were found in our own recent collections, records from recent literature. One important collectio n,

made by the 1965/66 Biological and Geological Expedition to Marion and Prince Edward Islands (Van Zinderen Bakker e/ a/. 1971). unfortunately could not be traced. This collection was the basis for Lindsay's (1977) account of the Marion and Prince Edward Island lichen nora. We have doubts about some of Lindsay's (1977) determinations, but unfortunately these doubts can not be substantiated by reference to the relevant specimens. We have decided not to include the following species from the latter account , none of which are

verified by other studies: Buellia coniops (Wallenb. ex Ach.) Th .Fr., Caloplaca vitellinella (Nyl. ex Cromb .) Zahlbr. , C/adonia balfourii Cromb. f. chlorophaeoides (Vain.) Evans , C/adonia cf. coccilera (L.) Willd ., C/adonia macilenta (L.) Hoffm., Cladonia pyxidata (L.) Hoffm. , Lecanora polytropa (Hoffm.) Rabe nh. , Lecidea aubertjj B. de Lesd. , Lecidea homa/otera Nyl. ex Cromb., Lec/dea ct. interrupta Darb., Pannaria hooken (Barr. ex Sm .) Nyl., Parmelia saxatilis (L.) Ach ., PamlOtrema cnnitum (Ach.) Choisy, Peltigera canina (L.) Willd. , Peltigera polydactyla (Neck.) Hoffm., Physcia cf. phaea (Tuck.) Thoms., Placopsis bicolor (Tuck.) B. de Lesd., Placopsis cf. cribellans (Nyl) Ras. , Ramalina cf. sublarinacea (Nyl. ex Cromb.) Nyl., and Xanthoria elegans (Link.) Th.Fr. Of 100 species listed in this account only 31 have been fo und on both islands. Most species (54 species) were only reco rded on Marion Island, while 13 species were only fou nd on P rince Edward Island. T hus the number of species found on Marion is about double that found on Prince Edward

Island (87 and 44 species respective ly). This may be explained by the fact that the time and effort expended in collecting on Marion has been much greater than on Prince

Edward Island . In addition , Marion is appreciably larger than Prince Edward Island , wh ich should also result in a larger

571

South Afflcan)ournal or Botany 2001, 67 552- 572

number of species in the former island . Because the climate and the geology of the islands is very similar, as well as the general vegetation stru cture, it is not likely that significant habitat~differences could explain the difference in species number, or the large number of species that have been found on one island only. From this we infer that on both islands the lichen flora is considerably richer than is presently known. Nevertheless it is strange that such a common lichen as Rhizoplaca melanophthalma is not found on the Prince Edward Islands. Also those lichens that appear to be ci rcuma ntarctic look somewhat different on these islands (see e.g. the discussion under Ca/oplaca isidioc/ada). Of the 100 taxa listed for the islands, six are new species. Most of these are from groups that are inconspicuous in the field. For this reason they may have escaped detection by previous collectors , most of which were not lichenologists. From our experiences at Marion and Prince Edward Islands, we expect that lichenological exploration of most other subAntarctic islands would yield a considerable number of new species as well. In addition we believe that such exploration will reduce the relatively large number of lichen species at present only known from Marion and Prince Edward Islands. The lichen nora of the islands of Ihe Southern Ocean is still very incompletely known , and many taxonomic problems still have to be resolved. This makes any delailed biogeographic account premature. Nevertheless, some patterns may be of significance. Of 95 lichens identified to species level , 31 also occur in the Northern Hemisphere. Twelve of these appear to be bipolar, while 19 species are more or less cosmopOlitan. Of the Southern Hemisphere species 25 are found exclusively to the easl of the islands (i.e. in Kerguelen, Crozet, Heard, Macquarie, Austra lia or New Zealand) . Only six species are recorded exclusively from areas to the west of the islands (South Georgia, Antarctic Peninsula, etc. ). Th is suggests a predominant dispersal pattern from east to west, against the presently dominant wind directions. This contrasts with findings by 0vstedal and Smith (2001), who concluded Ihat the lichen flora of South Georgia is mostly derived from regions to th e east of th e latter island, i.e. from southernmost South America. One-third (33) of the islands lichen species is presently nol know from areas outside the Kerguelen phytogeographical province. Fifteen of these have o nly been recorded from the Prince Edward Islands. It seems premature to decide that these are all endem ic, because the information on the prese nce especially of many inconspicuous lichen species o n most sub-Anta rctic islands and other Southern Hemisphere regions is very incomplete. As the Prince Edward islands geologically are quite young, their flora canno t be used in a discussion of Gondwanaland d istribu tions, as they have all immigrated long after the split-up of th e land masses. We are indebted to Or WAArcher, Canberra, Dr P Clerc, Geneve, Dr JA Elix, Canberra. Dr PM J0rgensen, Bergen, Dr H Mayrhofer, Graz, Mr RS Poulsen, Copenhagen. Dr U Srehling, Copenhagen, Dr T T0nsberg. Bergen and Dr 0 Vitikainen, Helsinki, for determination of or comments upon specimens, and to Mr J Serge and Mrs BH lngvartsen, both Bergen, for help with illustrations. and an anonymous reviewer for extensive comments. NJM Gremmen was partly supported by the Netherlands An tarctic Acknowledgements -

Programme. Logistic facilities were provided by the South African Department of Environmental Affairs and Tourism. We thank Prof. VR Smith, University of Stellenbasch, who initiated this work, far his continuing support. We are also indebted to Dr A Henssen, Marburg, for giving us access to her collections from Marion and Prince Edward Islands, and Dr J0rgensen for correction of the Latin diagnoses. References Archer AW (1997) The Lichen Genus Pertusaria in Australia. Bibliotheea lichenologica 69, 249 pp. Broda 1M (1972) The lichen Genus Coccotrema in North America The 8ryologist 76: 26G-270 Chown SL. Smith VR (1993) Climate change and the short-term impact or Fera l house mice at the sub·Antarctic Prince Edward Islands. Oecologia 96: 508-516 Coppins BJ (1987) The genus Ramonia in the British Isles. The Lichenologist 19: 409-418 Crombie JM (1877) Li chens Terrae Kergueleni - an enumeration of the lichens collected in Kerguelen Land by Rev. A.E. Eaton during the Venus-Transit Expedition in 1874-75. Journal of the Linnean Society (London) , Botany 15: 180-1 93 De Villiers AF (1976) Littoral ecology of Marion and Prince Edward Islands, (Southern Ocean). South African Journal of Antarctic Research . Supplement 1: 1-40 Dodge CW (1948) Lichens and lichen parasites . B,A.N.Z. Antarctic Research Expedition 1929-1931 reports , series B (zoology and botany). vol. VII 1-276 Egea JM, Torrente P (1994) EI genera de hongos Jiquenizados Lecanactis (Ascomycotina). Bibliotheca Lichenologica 54, 205 pp Follmann G (1968) FelsbewohnendeArthoniaceen der chilenischen PazifikkOste. Wildenowia 413: 365-382 FuHer NR (1967) A preliminary report on the littoral ecology of the Marion and Prince Edward Isla nds. South African Journal of Science 63: 248-253 Ganoway OJ (1985) Flora of New Zealand : lichens. Wellington , PO Hasselberg , 662 pp. Galloway OJ (1997) Studies on the lichen genus Sticta (Schreber) Ach. IV. New Zea land species. The Lichenologist 29: 105-168 Gremmen NJM (1981) The vegetation of the Subantarctic islands Marion and Prince Edward . Dr. W. Junk Publishers, The Hague. 149 pp. Gremmen NJM (1982) Alien vascular plants on Marion Island (subanlarclic). CNFRA 51 : 315-323 Gremmen -NJM, Smith VR (1999) New records of alien vascular plan ts from Marion and Prince Edward Island, sub-Antarctic. Polar Biology 21 . 401-409 Hall K (1978) Evidence for Quaternary glaCiation of Marion Island (sub-Antarctic) and some implications. In: van Zinderen Bakker EM (ed.) Antarctic glacial history and world palaeoenvironments. Balkema. Rotterdam , pp 137- 147 Henssen A (1963) The North American species of Massa/angia and generic relationships. Canadian Journal of Botany 41 :1331-1346 Henssen A (1969) Eine Studie Ober die Gallung Arctomia. Svensk Botanisk Tidskrift 63 : 126-138 Henssen A (1985a) Hertella, a new lichen genus in the Peltigerales from the southern hemisphere. Mycotaxon XXII (2): 381-397 Henssen A (1985b) A new Zahlbrucknerella from the subantarctic. The LichenoJogist 17: 301-303 Henssen A (1986) Edwardiella mirabilis, a holocarpus lichen from Marion Island. The Uchenologist 18(1): 51 - 56 Henssen A, Budel B (1984) Phyffisciefla, a New Genus of the Lichinaceae. Seihef! 79 zur Nova Hedwigia: 381 - 392 Henssen A, James PW (1982) Th e lichen genus Steinera. Bufietin of the British Museum of Natural History, Botany 10(4): 227-256

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