Polarforschung 58 (2/3): 65-76. /988

1.1 Problems in Monographing Antarctic Crustose Lichens

By Hannes Hertel*

Summary: OUf present knowlcdgc 01' thc taxonom)' nnd floristics of Antarctic crustosc lichcns is still pOOl' and unrcfiahlc. for thc following rcasons: I. lack 01' floras und monographs uscful for idcntification of antarctic Iichcns. 2. unavailability 01'imponant type collcctions. 3. lack of represemative collcctions. adequate 10 study variability and distribuuon. 4. undcrcstimation of thc cxtcnt 01'phcnotypic vmiation cuuscd by various abiotic anel biotic factors such as: phvsical f'actors (light. tcmpcruturc and moisturc conditions. freczc-thaw actions.wind erosion). ehernieal factors (growth sumation in nitrogen-cnrichcd niches. mincral incorporation in lichcns growing on substratas rieh in metals). animaIFeeding (in Antarctica by mites). fungul parasitism. andl.,a:~b2n,o~;n;:11:~a:l,'~F':~~;~:;~~~;H'S 5. unfurniliarity with known Irom other cool temperate in the world (Ieading to thc üc namcs). In an nppendix morc 100 liehen spccics rcportcd from Antarctica also occurring in othcr rcgions are

Zusammenfassung: Ursachen unserer derzeit geringen und wenig zuverlässigen systematischen und floristischen Kenntnisse antarktischer Krusten­ flechten sind: I. Mangel an Florenwerken und Monographien. die zum Bestimmen antarktischer Flechten hilfreich sein könnten. 2. Unzugänglichkeit wichtigerTypus-Aufsall1!11lungen. 3. Mangel an für das Studium VOll Variabilität und Verbreitung der Sippen 4. unzureichende Kenntnisse der hohen Modifikabilität dieser Flechten. SIC. physikalische Standortfaktoren (Bclcuchtungs-. Tempcra­ tur-, Fcuchtiakcitsvcrhältnissc. Wechselfrost. Windschliff). teils durch chemische Faktoren (Wachsturnsfördcrune an stickstoffreichen Standorten. Mineralein- und -auflugerungeu bei chalcophilen Flechten). tcils durch Tierfraß (in der Antarktis durch lv1ilben~ teils durch Befall mit pilzliehen Parasiten oder durch abnorme Regeneration bewirkt wird. S. mangelnde Kenntnis der aus" anderen kalttemperierten Regionen unserer Erde bereits bekannten Arten (\\"a~ zu zahlreichen überflüssigen Neubeschreibunzen geführt hat). In einem Anhang werden Über 100 nicht-endemische. aus der Antarktis gemeldete Flechtenarten gelistct. um den hohen Anteil weitverbreiteter Arten dieser Flora zu verdeutlichen.

1. INTRODUCTION

Scientists working on the terrestrial vegetation 01'Antarctica are usually confronted with a flora where lichens playamajor and often dominant rölc. When the c1imate becomes very harsh (as, e.g .. south 01' 75°S). fruticose anel foliose lichens elisappear anel saxicolous crustose species form the main component 01' the liehen vegetation (FRIEDMANN 1982).

Same 75 percent 01'the 13.500 species 01'lichens recognizee! at this time (HAWKSWORTH et al. 1983) are ..crustose lichens" [a biologicaL not a taxonomic group]. These smaillichens e!evelop only a thin, primitively organizee! thallus, which is either completely affixee! to the substratum over the whole lower surface (like a varnish, rarely thicker than 0.4 rnm), or is hie!den in the uppermost layer01'rock [0.2-1 [-4] mm in ene!olithic calciphi laus lichens (DOPPELBAUR 1959); up to 10 mm in Antarcitc ..cryprocndoliths" (FRIEDMANN 1982)].

Crustose lichens can colonize extreme nival ane!polar habirats. far better than other macroscopically visible plants. In Antarctica crustose lichens have been faune! as far south as 86°06'S (e.g. Lecidea cancrifonnis"; DODGE ane! BAKER 1938). in the Alps up to the highest icefree rocks (4810 m; Lecanora polvtropa; SCHLAGINTWEIT 1855). ane! in the Himalayas even up to 7400 m (Carbonea vorticosa and Lecanora polytropa: HERTEL 1977). None 01'the mountaineers who c1imb the 8000-meter peaks in the Himalayas have yet faune! time 01' interest to look far lichens at these high elevations. It is very likely that no altitue!inal ar southern climatic limit exists 1'01' crustose lichens on earrh. in contrast to bryophytes, foliose lichens. and vascular plants.

Bryophytes have been collecteel as far south as 84°37'S in Antarctica (Sarconeurnm g/acia/e: KAPPEN 1983). up to 4230 m in the Alps (Grimmia donniana. PITSCHMANN ane! REISIGL 1954. GEISSLER 1982). anel they grow at least at an elevation 01'6200 m in the Himalayas' (V KOMARKOVA. pers. comm.). Foliose lichens 01'

;' Prof. Dr. Hannes Hertel. Botanische Staatsammlung. Mcnzingcr Straße 67,8000 MÜnchen 19. Federnl Rcpuhlic 01'Germany =For ful! name (inc1uding aurhors namc) sec Appendix I. , A dense bryophyte vcgctation containing 8 spccics 401' hcpatics is rcported in ahitudes betwecn 5750 IIIand 6060 111. from hot spots on the Socompa Volcano on the Argentina-Chile (HALLOY 1986).

65 the genus Umbilicaria, have been collected as far south as 7T55'S in Antarctica (SIPLE 1938), up to 4640 m in the Alps (SCHLAGINTWEIT 1855), and up to 6300 m in the Himalayas (FREY 1936, POELT 1977). Vascular plants reach 68°21'S (Deschampsia antarctica, Anonymous 1985),4270 m (Ranunculus glacialis; PODHORSKY 1939) in the Alps and 6250 m (Arenaria musciformis; PODHORSKY 1939) in the Himalayas.

In spite of their outstanding röle as pioneers in extreme cold deserts, and in spite of the fact that these regions have a small flora relative to temperate regions, our knowledge of the taxonomy and distribution of Antarctic crustose lichens still is very poor and unreliable. It is even extremely difficult to give a realistic assessment of the number of lichens present in Antarctica (Antarctica, circumscribed here in the sense of WACE (1960, 1965), including the Antarctic Continent, the South Shetland Islands, the South Orkney Islands, the South Sandwich Islands, and Bouvet Island). According to DODGE (1974) there are 429 species oflichens (including 317 crustose ones) in Antarctica. This number is, however, as LAMB (1970) has already mentioned, undoubtedly excessive, and 160 species, in my opinion, is a more realistic estimate.

The reason for such a discrepancy - 429 versus 160 species - sterns from the low reliability of data published by some authors active in the flaristics and taxonomy of antarctic lichens. A good example of this situation is represented by a comparison of two recent revisions of Usnea subgen. Neuropogen (a conspicuous frutescent liehen genus, weil known to all terrestrial biologists working in Antarctica). In his liehen flora of the Antarctic Continent DODGE (1974) recognizes no less than 21 different species of Neuropogon in Antarctica. In WALKER's detailed monograph (1985), 17 of these 21 "species" are considered as synonyms. One of the remaining four species was excluded, being based on incorrect identifications (the species in question does not occur on the Antarctic Continent). Although WALKER accepts two additional species not included in DODGE's treatment, even so the total number of species for Antarctica is reduced from 23 to 5. While DODGE postulated all ofhis 21 species to be endemie to Antarctica, WALKER found that none ofthe accepted 5 species is restricted to Antarctica.

I am confident that this reduction in numbers of species, as shown here, can be generalized across all genera (see also "Appendix II", and it is entirely possible that some 80% of the 149 new species introduced by DODGE (1974), and DODGE and BAKER (1938) and accepted in DODGE's Flora (1974) will turn out to be synonyms.

I am convinced that it requires more experience and knowledge to understand the antarctic lichens than the lichens of any other region, and I believe the following factors account for the difficulties encountered in correctly identifying Antarctic crustose lichens.

2. DIFFICULTIES IN IDENTIFYING ANTARCTIC CRUSTOSE LICHENS 2.1. Lack 01 modern floras and monographs 2.1.1. F I 0 ras

Modern (post-1950) liehen floras which include relatively accurate keys to crustose lichens exist for only a few European regions: Western Europe (CLAUZADE and ROUX 1985), France (OZENDA and CLAUZADE 1970), S. W.Gerrnany (WIRTH 1980), Poland (NOWAK and TOBOLEWSKI 1975, NOWAK 1983), Ukraine (OXNER 1956, OXNER 1968). The liehen flora for the Soviet Union is not yet complete; five volumes published so far (KOPACZEVSKJA et al. 1971, OXNER 1974, BLUM et al. 1975, KOPACZEVSKAJA et al. 1977, GOLUB­ KOVA et al. 1978). Crustose lichens ofextra-European regions are very incompletely known, and for this reason treatments for the floras of Israel (GALUN 1970), Saudi Arabia (ABU-ZINADA et al. 1986), and New Zealand (GALLOWAY 1985) should be considered as first steps toward a better knowledge rather than a reliable means for identifying species of difficult genera such as Candelariella, Caloplaca, Lecanora or Lecidea.

DODGE's 'Lichen Flora of the Antarctic Continent and Adjacent Islands' (1974) is not a flora in the true sense where one can find help in identifying plants. REDON's (1985) account of the lichens ofthe Antarctic peninsular region ("Liquenes Antärcticos") is of relatively little help in deterrnining most crustose genera (except for Buellia and Verrucaria, where he follows LAMB 1948a, and LAMB 1968). Not a flora, but a valuable aid for identification of various groups of lichens are the keys by POELT (1969) and POELT and VEZDA (1977, 1981), although the large, difficult genera of crustose lichens (e.g. Acarospora,

66 Bacidia, Bue/lia, Cladonia, l.ercmora, Lecidea. vernuaria) are omitted or only partly included.

2.1.2. C h eck 1 ist S

Among various recent checklisrs, SANTESSON's (1984) critical. highly reliable and informative (also key to literature) list of the lichens of Sweden ancl Norway is of great value to all who are working on lichens of cool temporare ancl cold climates, both in the Norhern and Southern hernisphere, because ofthe high number ofbipolar and widespread lichen species in Scandinavia.

2.1.3. Mono g ra p h s

The number of monographs relevant to Antarctic lichens is still very smal1. There are worldwide monographs for Acarospora (MAGNUSSON 1929 - out ofdate), Cladonia (VAINIO 1887-1897 - out of date), Ochrolechia (VERSEGHY 1962 - out of date), Plucopsis (LAMB 1947). Spliaerophonts (OHLSSON 1974), Stercocaulon (LAMB 1977, LAMB 1978), ancl Usnca subgen. Neuropogon (WALKER 1985). For none ofthe large genera of erustose lichens (e.g. Buellia, Caloplaca. l.ccanora, Lecidea. . .) are there comparable monographs. Spccies of Buellia and Rinodina from the Antaretic Peninsula arc trcated by LAMB (1968): a prelirninary key for the saxicolous species ofLeeidea sensu ZAHLBRUCKNER in the subantarctic region (including a few species from Antarctica) is provided by HERTEL (1984).

2.2. Unavailabilitv oftvpe collections

Type materials of the more than 150 species of crustose antarctic lichens described by OOOGE and coworkers were not deposited in public herbaria, as indicated in publications, but kept in private herbann. For example, according to DODGE and BAKER (1938: 517), type specimens of 66 new lichens species described from Marie Byrd Land and King Edward VII Land should have been deposited in the herbarium of the Missouri Botanical Garden 01' in the Uni ted Stares National Herbarium in Washington (where the MO collection was latertransferred). In fact these specimens were incorporated in DOOGE's private collcction. which was recently transferred to Farlow Herbarium (FH) and is now finally available for loan and study.

A number of HUFs type specimens (HUE described more than 70 species of crustose lichens from Antarctica) seem to have been lost or misplaced (I was unable to trace these samples at Paris in 1987), although LAMB was able to locate so me in the 1950's.

2.3. Lack ofadequate collectious

Adistressing1y high percentage of lichens collected in Antarctica is sterile, badly developed, or damaged. Usually the specimens are very small (sometimes a few badly developed ascocarps only), much too small to tell anything about the variability of characters.

Tiny and inconspicuous species are extremly under-represented in collections from Antarctica, for many of the specimens wcre collected by geologists 01' amateurs.

Because of the general difficulties in determining Antarctic lichens, most of the existing liehen collections from Antarctica lie unstudied and undetermined. Therefore, monographers interested in the material of a particular genus have to visit the herbaria in queston to find and select specimens.

2.4. Underestimation ofthe extcnt ofphenotypic varianon

As SANTESSON (1952) has so apt1y pointed out, for a lang time many lichenologists practised a kind of taxonomy best described as .Jndividuum descriptions". Describing a single individual specimen or a small group ofspecimens does not take into account the normal variability and plasticity of any species, which may sometimes be extremely high. As POELT (1973) srates, phenotypes of a single species growing in different habitats can sometimes appear so unlike that it calls for detailed analysis and long experience to recognize them as representatives of one and the same taxon. However, increasing attention has been paid to variability and

67 modification in reeent years. WEBER (1962) ancl POELT (1973) have summarized the existing knowledgc. and some monographcrs have devotcd special chapters to the problern (e.g. HERTEL 1967, KILIAS 1981, SCHWAB 1986, WUNDER 1974).

Experience in arctic, alpine anclsubantarctic regions. as weIl as expcrienee from a careful study of large numbers of specimens from cool temperate and cold regions of the northern and southern hcmisphere, rather than field work in Antarcrica. has made me familiar with the high rate of obviously environmentally inducedmodifications in crustose lichens. Because of the more extreme and hostile environment in many parts of Antarcrica, an even lugher degree of modification should be cxpecred for lichens in Antarctica: FILSON (e.g. 1982) has drawn attention to this problern.

A nurnber of abiotic and biotic factors are responsible for this high variability, as pointec1 out by WEBER (1962) ancl POELT (1973),

Before going into details, it should be mentioned that in various cases ofobviously cnvironmcntally altered liehen specimens, we are not able to explain which panicular factors have caused the alteration. Often a cornbination of factors acting simultaneously or successivcly may be responsible for the modification in qucstion (see also WUNDER 1974).

2.4. l. P h y s i c 'I 1 fa c tor s . e, g.: 1i g h t . t e m per a t u re. m 0 ist u rc, fr e c z e - t h a w a c t i o n , wind erosion

Shaded andlight-exposed thalli often differ considerably in thallus colour, Light exposed parts of the thallus are usually more intensely colonred.

Specimens of certain spccies (e.g, Leeidee lapicidai colleeted at high elevation in the Alps 01' in corresponding parts of the Arctic usually devclop 'I much rnore conspicuous black prothallus than specimens growing under more temperate conditions do or tend to develop a thallus with isolared areolae or small groups of areoles on a more or less conrinuous black prothallus (as e.g. in some specimens of Rhizocarpon superficialei. [The ability to form a black prothallus seerns to be 'I genetically controlled character, but the degree of manifestation of this structure is environmentally conrrolled.] One might speculate that the black surface of the prorhallus will absorb more solar radiation and so elevate thallus tempcrature and (as long as water is available) assimilation rate.

While such conspicuous ancl enlarged prothalli are sometimes found in crustose species collected in the more humid maritime parts of Antarctica, this phenomenon secms to be rarer in dry continental parts of Antarctica. The majority of species there show well-developed, thicker epilithic or even substipitate or stipitate thalli (HALE 1987), My observations however, are so far based only on herbarium specimens (and not on population studies in the field). Interpretation of specimens in the laboratory is often distorred by alterations caused by wind erosion ('I specimen which scems to c1eveloponly traces of an epilithic thallus may, in reality, produce a well-developed epilithic thallus which subsequently becomes more 01' less totally eroded). Population studies in the field, or at least much richer collections than are presently available, are neccessary to understand the extent of this kind of variability. It is difficult to decide whether a species such as Car!Jol1ea 1'orlie05a, which usually has a strongly reduced epilithic thallus when growing in alpine and nival belts ofthe Alps, Himalayas or in the Arctic, developes a distinct stipitate epilithic thallus when growing in the cleserts of Antarctica, or whether two closely related but genetically distinct species (Carbollea \'Orticosa, Cal'bollea capslI!ala) are involved. This is the kind of problem which is extremely difficult to answer in lichenology.

Freeze-thaw action probably favours pulvinate thallus surface, als FILSON (1982) suggested.

Especially in regions such as continental Antarctica or the Himalaya above 7000 m altitude (KUHLE 1986), where snow permanently clrifts around (because the temperature necessary for f01'l11ing ice or firn cannot be reacbed), very slow-growing crustose lichens suffer severely from the strong abrasive action of snow crystals and sand. Unusual alterations in gross morphological characters are therefore common in continental Antarctica,

68 2.4.2. C he m i c a I fa c tor s

As WEBER (1962) points out. stimulation of growth is wcll known in lichens from nitrogen-enriched sites (e.g. ornithocoprophilous lichens). Crustose lichens are more luxuriant and usually develop thick, sometimcs almest squamulose thalli. However. only a small number of species in genera such as Buellia, Candclariclla. Caloplaca. Lecanora, Lccidella. and Rinodina tolerate high nitrogen enrichment. In cases where maeroscopie thallus characters appear to have taxonomic significance, luxuriant modifications may be troublesomc. An example is Lecidella inamoena. normally an enelolithic speeies. In the Arctic this taxen was elescribed as .Lecidella endoli-thea' (the enelolithic growth form) anel as .Lecidea spitsbergensis" (a specimen with a well devcloped epilithic thallus). Although it is not mentioneel on the Iabel that Leeidea spitsbergensis was collected near a seabird colony (unfortunately ecological data are rarely giveu on specimens labels), I am almost certain that these two taxa are no more than ecotypes ofone anel the sarne species. However, it is still not clear whether the very similar eu-thalline Leeidee alaiensis. elescribeel from the Alai mountains in central Asia, is a disrinct taxon or not.

It is well known that sorne crustose lichens telerate high levels ofheavy metals toxic to most organisms, Leea-nara vinetorum has a copper content of 0.5% of the dry weight of the liehen (POELT and HUNECK 1968); Leeidee lapicida (= Leeidea theiodes) up to 5.3% Cu dry wt. (PURVIS 1984); Acarospora sinopica up to 6.5% Fe elry wt (LANGE anel ZIEGLER 1963). Lecidca theiodes is, as PURVIS (1984) points out, a modifieation of Lecidea lactea (which is, as suggested by SCHWAB (1986). a chemotype of Lecidea Iapicidai. The characteristic anomalous green coloration of the thallus of Lecidea theiodes, which led SOMMERFELT to describe a new species, is caused by crystals of a eopper-norstictic acid complex. as it is in two other yellow-green species elescribed from cupriferous rocks: Acarospora isortoqucnsis, and Acarospora undata, wh ich were found to be modifications of Acarospora smaragdula (PURVIS et a1. 1985. PURVIS et al. 1987).

Lichens growing on weathered, iron-rieh rocks often show a rusty coloration, caused most likely by iron oxide. As for the lecideoid lichens, SCHWAB (1986) has rccently shown that aside from a number of species with constantly rust-coloured thalli (e.g. Tremolccia atratai. most rust-coloured lichens described are modifications of whitish 01' grcyish species (this was suspected by various lichcnologists. e.g. WEBER 1962).

Ochraceous pigmented moelifications seem to be restriored to cool anel humid climates, and thus so they are very common in the subantarctic islands, known also from maritime Antarctica, but not yet from Continental Antarctica.

2.4.3. Bio t i c fa c tor s , s u c h a san i mal fee d i n g an d fun g alp a ra s i t i s m

In ternperate climates. damage to thalli and/or ascocarps caused by predation by small animals, e.g. caterpillars (RAMBOLD 1985), snails (SCHADE 1933, SCHADE 1956). colembolans and mites (GERSON anel SEAWARD 1977), plays a significant role in the moelification of lichens. In many especially eutrophie habitats of temperate Europe, for example, it is sometimes quite elifficult to find any undamaged thalli ofcertain speeies (POELT 1973). We usually elo not know what kind of animal causeel the darnage. except in freshly damageel thalli where, for example, radula tracks on cortex and medulla of the liehen can be ciearly observed. Gross morphological characters (type of thallus areolation, shape of ascoearps) may be strongly altered.

Hymenia, structures rich in protein, often become selectively eaten. In lichens with carbonizeel excipular strucrures (as in species of e.g. Bue/lia, Car!Jonea. Carillaria. Porpidia, Rhizocarpon) this type of dal11age l11ay lead to apothecia which seem to have eleveloped very prominent anel often narrow margins anel flat 01' concave eliscs.

Another example of a misunderstood feeding elamage form is ,,Lecida c)'closora". a lichen characterized by roundish soredia, which are in reality the basal parts of the apothecia of PO/pidia zeoroides, totally eaten away (HERTEL 1967). I have had no personal experience on whether animal predation of lichens plays a significant role in the l110dification of crustose lichens in Antarctica. However. LAMB (1968) found "elamage by mites (AcarinicJae) with subsequent abnormalities in regeneration" .significant for the understanding of polymorphie variation in species of Buellia in the Antaretic Peninsula. Most high latitude parts of Antarctica are too colel to support any invertebrates.

69 Worldwide there arc some 500 species offungal parasites on Iichens, belonging mainly to the Ascomyeotina and Imperfect Fungi, Some fungal infections eause descolorations or gall-like structures. and massive infeetions may suppress development of ascocarps or ascospores of the host liehen. Ascocarps of the parasite grow on a sterile thallus and the whole system then is often misunderstood as a "normal liehen". In this way HUE (1915) described Charcotia as a new genus of Umbilicariaceae in Antarctica, based on material of Umbilicaria antarctica parasitized by a species of Scutula (LAMB 1948b). 111e"antarctic genus Lethariopsis" is based on some rather old speeimens of Usnea antarctica with a few minute fruiting individuals of a Caloplaca species epiphytically attached to them (LAMB 1948b). Similar misinterpretations will probably be found.

Endohymenial parasites are sometimes reduced to ascogenous hyphae and asci, which occupay the aseocarps of the host, surrounded by the host paraphyses. Combinations of asci and spores of the hymenial parasite Arthonia intexta and the ascocarps and thalli ofthe hostLecidella have been described nine times as species new to science (HERTEL 1969).

Little is yet known of liehen parasites and their mcdifying influence on host lichens in Antarctica.

2.4.4. Ab n 0 r mal re gen e rat ion

The ability to regenerate damaged structures is well developed in crustose lichens. However, the resulting regenerated thalli or ascoearps may differ considerably in morphology frorn the undamaged species.

Carbonized structures may develop around damaged tissues. An example is the subantaretic liehen Lecidea lygomma, wh ich forms remarkable blaek rings around the apotheeia (HERTEL 1984). This charaeter was so eonstant throughout the large populations I studied on the Prince Edward Islands that I became convineed it was a good taxonomie eharaeter. I used it to establish a new genus Zosterodiscus (HERTEL 1984). However, after later studying New Zealand populations, I concluded that this eharacter is not constant and therefore the carbonized rings must be environmentally, not genetically induced structures (HERTEL 1987b).

Regeneration of hymenia may lead to a gyrodisc (HERTEL 1967, KILIAS 1981) and umbonate ascocarp.

However, not only gross morphological, but also intemal anatomieal eharaeters may be altered by damage and subsequent regeneration. Normally greenish or green-blaek epihymenia may become brown (SCHWAB 1986). In regenerated hymenia there may be a greater degree of branching and anastomosing of paraphyses, usually 'I good taxonomie eharacter at the genus level, than in undamaged ones (KILIAS 1981, SCHWAB 1986). SCHWAB (1986 p. 278) mentions that the width of exeipular hyphae (usually a good taxonomie eharaeter) may also be altered by darnage and regeneration in Porpidia.

Development as well as regeneration are very slow processes in Antaretica. So a new wave of damage often may occur before an earlier damage is regenerated. In this way very strange thallus and ascocarp struetures may develop.

It is usually easy for plant collectors to deeide whether a flowering plant is well developed, an extreme shade form, variously damaged (e. g, leaves damaged by insects, or stunted by fungal parasites), without flowers or fruits, or represents the remains of aplant eaten by grazing animals. In all these cases a good trained collector would do his best to find additional, better developed speeimens.

Analogous kinds of damage obviously oecur in lichens. However, in these cases it is often diffieult to recognize whether there is any damage at all and what the nature of it is,

The extreme variability of morphological characters, eombined with our present inability to eulture these extremly slow-growing, saxieolous, crustose lichens, makes it very difficult to deeide whether two similar looking specimens are either merely eeotypes of one and the same taxon or are genetically different.

70 2.5. Lack offamiliarity with Arctic and Alpine lichens

The degree of endemism among Antarctic crustose lichens in apparently very much overestimated. According to DODGE (1974), nearly all (97%) of the lichens found in Antarctica are claimed to be endemie. I consider 20% endemie species to be a much more realistic estimate. The reminder of the species have abipolar 01' otherwise wide distribution. In "Appendix H", I have tried to summarize the non-endemie species in the Antarctic liehen flora, as far as I could determine (the list, based on literature available to me, is undoubtedly incomplete). There are 69 species which may be increase when genera such as Aearospora, Bacida, Buellia, Catillaria, Caloplaca, Lecanora, Leeidea, Rhizocarpon and others are monographed on a geographically wide base. Lichenological research in Antarctica started very late, and most of bipolar or widespread species were described much earlier, often in the 18th and 19th centuries, from the N0I1hem Hemisphere, mainly from Central Europe and Scandinavia. Therefore, a profound and detailed knowledge of these floras is an absolute prerequisite when monographing Antarctic crustose lichens 01' describing new species.

It is usually taken for granted that there is a sharp contrast between the liehen floras of the humid and perhumid subantarctic regions (Tasmania, New Zealand, W. Patagonia, Falkland Islands, South Georgia, Tristan da Cunha, Gough, Prince Edward Islands, Crozet Island, Kerguelen, Macquarie Island), and Antarctica. Appendix H lists 38 southern hemisphere liehen species which share a distribution in both maritime Antarctica (South Orkneys, South Shetlands, Antarctic Peninsula) and subant-arctic regions. This is not particularly surprising, for in subantartic regions there may be habitats (such as at higher altitudes on mountains) where microclimatic situations correspond to macroclimatic conditions in maritime Antarctica.

Although Antarctica is geologically very old, and was formerly apart of Gond wanaland, the opportunities for survival of Gondwanan floral elements are extremely limited due to the heavy glaciation which began at least 20 million years BP (RAVEN and AXELROD 1974). In the lecideoid lichens there seem to be no genera truly endemie to the Antarctic continent, in contrast to subantarctic regions (HERTEL 1987a).

There is no doubt that there are endemie crustose species in continental Antarctica (among the lecideoid species, Leeidea cancriformis, widespread and common in continental Antarctica, seems to be a good example). However, as in the Arctic, endemie species are often a minority among the more aggressive, widespread orcophytes, which recolonized the polar areas after Pleistocene glaciations by means of long distance dispers al.

One ofthe fascinating goals of liehen systematics is to clarify the origin of the Antarctic liehen flora, its diversity and relationships. We urgently need reliable identifications, and specialists who not only study Antarctic lichens exclusively but are also familiar with all other species described from various cold areas. Last but not least, we need more extensive collections from all ice-free parts of Antarctica made by trained lichenologists.

3. ACKNOWLEDGEMENTS

I wish to express my sincere thanks to a number of colleagues who provided me with various informations or who made available rare literature: Dr. Vera KOMARKOVA (Boulder, Colorado), Dr. O. BREUSS (Wien), Prof. Dr. W. FREY (Berlin), MI'. P. JACOBSEN (Kiel), Prof. Dr. L. KAPPEN (Kiel), MI'. G. RAMBOLD (München), D1'. R. LEWIS SMITH (Cambridge), Prof. Dr. J. POELT (Graz), MI'. F. SCHUHWERK (Regensburg), and Dr. O. VITIKAINEN (Helsinki). Dr. D. GALLOWAY (London) and Dr. M. E. HALEjr, (Washington) kindly revised the English text.

71 Appendix I

Index of Latin names mentioned in the text

Acarospora Massai. 67 Acarospora isonoqucnsis Alstrup · 69 Acarospnra sinopica (Wahlcnh.) Kocrb. 69 Acarospora sntaragdula (Wahlcnb.) Massal. 69 Acaraspora undata Clauzadc. Roux ct Wirth 69 Arenario tnuscifannis Wall. .66 Arthonia intexta Almqu. .70 Bnctlia DNol. . 66.68.69 Candelarictta Müll.Arg. 67 Carbonca capsulata (Dodgc) Haie 68 Carbonca vonicasa (Flk.) Hertel . 65.68 Charcotia Hue · 70 Ctodina Nyl. · 67 Cia donia Hill. cx Web. in \Vi211. · 67 Desclunnpsia antarctica Des,,~-­ 66 Grinnnia donniana Sm. 65 Lccanora polytropo(Hoffrn.) Rabenh. · 65 Lccanora vinctonnn Poet! ct Huneck .69 Lecidca ataicnsis Vain. 69.7() Leeiden cancnfonnis Dodgc ct Bakcr 65.71 Leeiden cvclosora Lettau 69 Lccidea cndolithco LVIH!e 69 Lccidca lactea Schac~. -- · 69 Leeiden Iapicida (Ach.) Ach. . 68.69 Leeiden lyrzOinmaNyl. · 70 Lecidca senst! Zahlbr. · 67 1J:;';;:;:':;"~~;;;:l:~J!. · 69 Lecideal.ccidca tJj .69 Lecidcita inamoena (Müll.Are..) Hertel .69 Lethanopsis Zahlbr. ~ · 70 Ochrolcchia Massai. 67 Placopsis gelida (Nyl.) Linds. · 67 POJ7Jidia zeoroides (Anzi ) Knoph ct Hertel .69 Ranuncnlns gtacialis L. · 66 Rhizocarpon snperficialc (Schacr.} Vnin. · 68 Rinodina (Ach.) S. F.Gray 67.68 Sarconcnntm glaciale (C. Müll.I CarcI. cl Bryhn 65 Scutula Tut. · 70 Sphaerophotus Pers. .67 Stereocanion Hoffm. .67 Tremolccia atrata (Ach.) Hertel 69 Umhilicaria antarctica Frcy et lVI. Lamb · 65 Usnea antarctica DR. · 70 Usnca P. Brownc cx Adans. subgcn. Neuronagon (Nccs ct Flot.) Jatta · 66 Zosterodiscus Hertel . .70

Appendix 11

This list ineludes a large number of non-endemie liehen speeies which oeeur in Antaretiea and in other parts of the world, Although most of the information is based on literature only, only those records are included, whieh scem to mc to be reliable. Usually only the more reeent literature eitations are included.

A. Bipolar and widespreadliehen species in Antaretiea

J. Acarosporachlorophana (Wahlcnb. in Ach.) Massal. (0VSTEDAL 1983) nigricans (Ach.) Ny!. (LAMB 1964. SMITH 1972, ALLlSON and srvJITH 1973) 3. Bacidia trachona (Ach.) Lettau (0VSTEDAL I986a) 4. Bryocauton chalvheifonnis (Ach.) Kärnef [= Alectosia chalvbcifonnis Ach.) (LAj'vIB 1964. SMITH 1972. ALLISON ct SMITH 1973) 5. Bucllia coniops (WahJenb. in Ach.) Th. Fr. (LAMB 1968, SlvIITH 1972) 6. Bucllia papitlara (Sommcrf.) Tuck. (LAMB 1968) 7. Buellia punctata (Hoffm.) Massal. (LAlvIB 1968. Si\'rITH 1972) 8. Caioplaca citnna (Haffm.) Th. Fr. [= Ca/oplaca mawsonii (Dodge) Dodge] (FILSON 1974a and 1974b. KASHIWADANI 1979) 9. Caloptaca jungcnnanniae (Vahl) TI1.Fr. (0VSTEDAL 1986a) 10. Catoplaca tetraspora (NyI.) Oliv. (1. POEL"L pers. comm.l 11. Caloplaca tiroticnsis Zahlbr. {J. POELT, pers. comm.} 12. Candcknielta vitellina (Hoffm.) Müll. Ar2. (BOWRA el al. 1966. SMITH 1972) 13. Carbonca vorticosa (Flk.) Hertel {= Leci(/ca vorticosa (Flk.) Koerb.j (HERTEL 1984, HERTEL 1985') 14. Cladina mitis (Sandst.) Hustich [= Cladonia nntis Sandst.. = Cladonia lacvignta sensu DODGE 1974] (AHTI and KASHIWADANI 1984) 15. Cladina rangifcrina (L.) Ny\. [= Cktdonia rangifcsina (L.) Nyl.. = Cladonia vicaria R. Sant.] (AHTI 196 L SMITH 1972. AHTI 1984, HUNECK et al. 1986)

72 16. Cladoniofurcata (Huds.) Schrad. (LINDSAY 1975') 17. Cladonio grocilis (L.) Willd. [:=Cladonia propogulifera (Vaiuio) Dcdge] (AHTI 1980. HUNECK el al. 1986) 18. Cladonio plcurota (Rk.) Schacr. (VAINIO 1903, LINDSA'{ ]975) 19. Cladonia pvvidara (L.) Hoffm. (VAINIO 1903. DARBISHlRE 1911. LINDSAY 1975. J()RGENSEN 1936) 20. Coclocoulon acnleattnn (Schreb.) Link (Schreh.) Ach.] (LAiVlB 1964. Sl'vfJTH 1972. KARNEFELT 1986) 21. COCIOCOllloll nnuicaunn (Ach.) Kämcf. llIuricara (Ach.) Ach.] (K/\RNEFELT 1986) 22. Cvstocoleus cbcncus (Dillw.l Thwaites (LINDSAY 1971b. SMITH 1972. SrvliTH and CORNER 1973. J0RGENSEN . 19X6) 23. Dennatocarpon intcstuufonne fKocrb.} Hasse (LAivIB 1948a) 24. Fansoldia dissifJahilis (Ny!.) I-ICrlel[= Lcridca slIhllllCSCCIIS NyL] (HERTEL 1984) 25. Hvpogvntnia IlIgubris (Pers.) Krog (LlNDSAY 1974. EUX 1979. REDON 1985) 26. l.ccanora potvtropa (I--loffm.) Rabcnh. (DARBlSHIRE 1912. LONGTON ct HOLDGATE 1979) 27. Lecidca atrobrnnnea (Ram.) Schacr. [= Lccidca racovitzac Dodge] (I-1ERTEL 1984) 28. Lecirtca lapicrda (Ach.) Ach. f= Lccidca ntpicida Vain.] (HERTEL 1984. HERTEL ]985) 29. Lecidello carpatlnca Koerb. (HERTEL incd.} 30. Lccidctto inamoena (Mtill. Arg.) Hertel (HERTEL ined.) 31. Lccidelio stigmatca (Ach.) Hertel and Lcuckert (HERTEL ined.) 32. Lenraria ncglccta Vain. (KAPPEN 19851 33. Mossatonoiacarnosa (Dicks.) Kocrb. (LAMB 1959. UNDSAY 1975, J0RGENSEN 1986 34. Ochrolrchiajrigida (Sw.) Lyngc (LlNDSAY 1971c, SMITH 1972. sxtrn: and CORNER 197J. REDON 1985) 35. Octnolrchia pa relkt (L.) Massal. l:= Ochrotcctua cnnarctica (Mull. Arg.) Darb.] (SivIITH 1972. 0VSTEDAL 1986b) 36. Pannaria hooked (Borr. es: Sm.) Nvl. (LINDSAY 1974. REDON 1985, J0RGENSEN 1986) 37. Pannclia saxantis (L.) Ach. [:=Parinclia ac('rmta Hue] rsxirru 1972. UNDSAY 1973a. StvnTH and CORNER 1973, UNOSA)' 1975) 38. Peltigem dtdactyia (Wirh.) Laundon l- Pettigcra spuria (Ach.) OC] (UNDSAY 1971a, S.ivliTH and CORNER 1973. UNDSAY 1975, VITIKAINEN 1987 in lit.) 39. Pettigere nqescens (Weis) Humb. ! 917a. SivlI1'H 1972. ALLISON er SMlTI-I 1973. LINDSAY ]975. VITIKA1NEN 1987 in lil.) 40. Phaconhi:a nintbosa (Fr.) Mayrh. c! Poelt [= Rinodina nimbosa (Fr.) Th. Fr.J (LAtvlB 1968) 41. Phvsciacacsio (Hoffrn.) Fürnrohr [:=PW"IIIe!lo("orcyiOoelge ct Bakcr.> Pannctiajolinstonii Dodgc. = Partnetto variolosa Dodge et Bakcr] (SMI1'H 1972. FILSON 1974a. NAKANISHI and KASH1\VADANI ]976) 42. dubia (Horfm.j Leuau (NAKAN1SHI and KASHIWADANI 1976, J0RGENSEN 1986 43. muscigena (Ach.) Poch (S1\·1I1'H 1972. KAPPEN ]985, REOON 1985) 44. Psendephebc minuscu!a (Arnold} Brodo ct Hawksw. f= /vtcctona tnnntscu!a (Arnold) Dcgcl.i e Atcctotia antarctica Dodgc ct BakcL:= Atcctotia bifonnis (Vain.I Dodge.> Alcctona cangesta (Zahlbr.) Dodge] (LAivlB 1964. KASHIWADANI 1970. BRODO and HAWKSWORTI-j 1977) 45. Psettdcphcbc pubcsccns (L.) Brodo et Hawksw. [ = Alcctoria pubcscens (L.) R. Howe. := Atcctona intricata Huc. :=: Atectona lIigcrrima Hue ] (LAMB 1964, HAWKSWORTII 1972. SMITH 1972) 46. PSOJ"OlIW hYPllol"IIJII (Vahl) S. F. Gray l- Psorotna tnvantii Dodge. = Psoroma ialhnannii Dodge] (SMITH 1972, HENSSEN Cl RENNER 1981. J0RGENSEN 1986) 47. Rhizocarpon gcographiCil/1I (L.) Oe. rsxnru 1972. ALLISON er Si'vIITI-l1973. Sj'vIlTH ct CORNER 1973. REDON 1985, HUNECKct al. 1986. 0VSTEDAL 1986a. 0VSTEDAL 19X6b) 48. Rhizocarpon supcrjicralc (Schaer.) Vain. [= Rliisocarpon adarcnsc (Darb.) ;V1. LAj'vlB.:= Rhizocarpon barilochcnsc Ras .. = Bucllia austro-gcorgica MUll. Arg.,:= Buclliaflavoplona Darb.. = Buctlia supcrbo Darb.. = Bncttia tristis Darb.. = Buellia variabilis Darb.] (RUNEi'vIARK 1956. SMITH 1972) 49. Rltizoplaca mclanoplultaltna (DC) Leuck. ct Poclt l- Omphalodina exulans (Th. Fr.) Dodge] (FILSON 1974b, FILSON 1975c, 0VSTEDAL 1986b) 50. Rmodina n!iracCOhl"!lIIlIm Doelge el Baker [= Rillodil/a arc!wcoides H. i\'1agn.] (LAiVlB 1968. KASHIWADANlI970. F!LSON 1975a) 51. RillOdillo tw:täcca(Wahlenb.) Koerb. (LAMB 1968, 0VSTEOAL 1986b) 51. Splwerophoms g{oboslIs (Huels.) Vain. (LlNDSAY 1971a. Sj\lITH 1972. SMlTH anel CORNER 1973, HUNECK er a1. 1986) 53. Spnras!afia tesflldillca (Ach.) MassaI. (HERTEL ined.) 54. Sferco('{/lIlolI alp,-mllli Lau!".(SivIITH 1972. ALLISON ct SMITH J 973. LAi'VIB1977) 55. Slcrcocalilol/ \"('S/f\'iallll/JIPers. (LAi\tIB 1977 - 79' 11'S~) 56. Tephmme/a atm (I-Iuels.) Hafellner Lecallora a/ra (Huels.) Ach.] (VA1Nl0 1903, SivliTH 1972, ALLlSON Cl Si\1ITI-l 1973) 57. Trel/lolicia atrata (Ach.) Hertel [= i:mCt.] (SivIiTH 1972. SMITH allel CORNER 1973. HERTEL 1984. HERI'EL 58. UlI1hilicaria aprilla Ny!. [= UlI1hilicaria spol1giosa Dodge el Baker, Umhilicaria all/arclica var. suh\'iridis Frey et M. LA!\·IB. UlIIlnlrcoriasm·i,·,i, Llano. Gyrophora kOi"Olkcl'ic:ii Golubkova] (GOLUBKOVA anel SAVICZ 1966, LINDSAY 197Ib, FlLSON 1975b, FlLSON 198/) 59. UlIlhilicaria dccllssar(l (ViiI.) Zahlbr. [= Ulllbi!icaria c-rill1ia Hue, Umhilicaria pal"l"lIla Huc. Umbilicaria mgosa Doclge ct Baker, UlI1hilicaria ccrchr~t"ormis Dodgc et Bake!". Ulllhilicana patenfol"lnis Doclge et Baker. Ulllhilicaria hU!1fcri Dodgc. Umbiticaria suhcachrtt"orlllis Dodgc. OJllplwlodisc/ls dcc/lssar/is var. !or!HOSIIS Llano. Ulllhiticaria Iciocmpa val'. nana Vainio. Dcrllla!isclIfllllwlI"So/li Oodge (LINDSAY 1969. KASHIWADANI 1970, SMITH 1972, FILSON 1975c. HUNECK cl al. 1986. FILSON 1987) 60. Ulllhilicariapropa/;lIlffcra Llano (TOPHAl--il el a1. SEAWARD el a1. 1983. FlLSON 1987) 61. USl1ca sphoce/(l/o R. Br. USllca frigida Dodge cl Baker. = laxissilllo Dodge. = USI1CO picala (lvI. Lamb) Dodge. := USJ/C(l scahridilla (1....1.Lamb) Doelge. = slllplw}"c(l Th. FI:,?:= [/.1'11('(1 s/riala Znmmulo in Oodge, = USllca suhpolaris (lvI. Lamb) Dodge] (LA MB 1964. WALKER 1985) 62. \'crl"/rcaria ccuthocmpa Wahlenb. (LAlVIB 1948a, SMITH ]972) 63. \icrrltcaria !II(fHI"(l Wahlcnb. (VAINIO 1903, Si\-HTH 64. \hnrcaria microspo/"o aucl. non Nyl. (LAi\-'lB 1948a sub microspora Ny!.'. SAN1'ESSON 1984) 65. Fcrl"/rcaria 1/Iucosa Wahlenb. (SANTESSON ]938. LAivIB 1948a. 0VSTEDAL 1986b) 66. Xal1t!lol"i(lUlndetwia (L.) Th. Fr, [= Polycaliliolla com{{igera Hue. Xalllhoria alllorelica (Vain.) Dodge cl Baker] (LAMB 1948b. SMlTH 1971. LlNDSAY 1975. J0RGENSEN 1986) 67. Xamhoria clegall.\" (Link) Th. Fr. [= Gasparrillia {{//slrogeorgica Dodge, = Gaspani/lia cirrina Dodge, := Gasparrillia harrisso/lii Dodge, := Polycaulionia johllstol/ii Oodge, = Polycalllio/lia pu!l'iJ/ata Dodgc et Bakc!"' := Po{ycou!ionia .\parsa Dodge et Bake!"' :=Ca!op{aca .\J)({IX[ (Oodge et Baker) J. ivfUlTayvar.latespora J. Murray] (FlLSON 1966, LINDSAY 1972b, Si'vlITH 1972. UNDSAY 1975. FILS ON 1975c. LlNDSAY 1977. FILSON 1984) 68. Xallt!wria parieti!w (L.) Behr. (KASHIWADANI 1979)

B. Species known!rom bOlh AllIarclica (md SuballIarclic regions

69. Bacidia tubaculma Darb. (0VSTEDAL 1986b) 70. BlIe{!ia anisolJlcra Vain. (LAMB 1968, LINDSAY 1973b. 0VSTEDAL 1986) 71. BlIet!ia augllsta Vain. (LA.1\·18 1968. LINDSAY 1973b) 71. BlIclliafa{klandica Darb. (DARBISHIRE 1912. 0VSTEOAL 1986b) 73. Bllc!lia isa!Jcllilla (Hlle) Darb. (LAMB 1968, LINOSAY 1973b) 74. Bucllia {atc!l1argillata Darb. (LA MB 1968. UNOSAY 1973b) 75. BHcllia IlIc{anosto{a (Hlle) Darb. (LAMB 1968. LINDSA\{ 1973b, 0VSTEDAI. 1986b) 76. Bucllia i"IISSO (Hue) Darb. (LAMB 1968. SivIITH 1972, LINDSAY 1973b, 0VSTEDAL 1986b) 77. CalojJ/a("(f cirrochmoidcs (Vain.) Zahlbr. (SMITH 1972. LINDSAY 1976. 0VSTEDAL 1986b) 78. CalojJ{aca rcgalÜ (Vain.) Zahlbr. (SMITH 1972, POEL1' ct PELLETER 1984. HUNECK ct aL 1986) 79. CalojJ{(f("(fsublohl/lata (Ny!.) Zahlbr. (SAN1'ESSON 1944. SivHTH ]971, REDON 1985. 0VSTEDAL 1986)

73 80. Cladina pycnoclada (Pcrs.) Leight. [= Cfadonia pycnoclada (Pcrs.) Ny!.] (HUNECK er al. 1986) 8 I. Coelocauton cpiphorcttunt (Ny!.) Kärnef (LAMB 1964, SMITH 1972. KARNEFELT 1986) 82. Haematonnna erythronnna (Nyl.) Zahlbr. [=Lepraria pallidostrcnninec Vain.. =Lcccnoraorosthcoides Vain.] (FOLLMANN and REDON 1971, SMITH 1972. SMITH and CORNER 1973. FOLLMANN and RUDOLPH 1979. HUNECK et al. 1986) 83. Hinumtormia lugulnis (Hue) M. Lamb (LAMB 1964, SMITH 1972. LINDSAY 1975, HUNECK et al. 1986) 84. Huea austroshctlandica (Zahlbr.) Dodge (LlNDSAY 1974) 85. Lecania gerlachei (Vain.} Darb. (0VSTEDAL 1986b) 86. l.eptogium pubcrulnm Hue (SMITH 1972, LlNDSAY 1975, REDON 1985, 10RGENSEN 1986) 87. Notolecidea subcontinua (Nyl.) Hertel [= Leeidee subcontinun Nyl.] (HERTEL 1987a) 88. Pannaria dichroa (Hook. et Tay!.) Cromb. (10RGENSEN 1986) 89. Parmelia ger/achei Zahlbr. (LINDSAY I973a. LINDSAY 1975) 90. Pannelia usliuaiensis Zahlbr. (LINDSAY I973a, LlNDSAY 1975) 9 I. Placopsis contonuplicata M. Lamb (LAMB 1947, SMITH ! 972, 0VSTEDAL 1987b) 92. Psoroma cinncnnomcnm Malme (LINDSAY 1974. REDON 1985) 93. Psoroma tcnuc Henssen (HENSSEN ct RENNER 1981) 94. Ramalina terehrata Hook. f. et Tay!. (LAMB 1964. SMITH 1972. LINDSAY 1975, REDON 1985, HUNECK et a!. 1986) 95. Rinodina dcccptionis M. Lamb (LAMB 1968, LINDSAY 1973b, 0VSTEDAL I986b) 96. Rinodinapctennannii (Hue) Darb. (LAMB 1968. SMITH 1972. LINDSAY 1973b) 97. Staurothole gelida (Hook. f. et Tay!.) M. Lamb (LAMB 1948a, REDON 1985) 98. Stercocaulan antarcticum Vain. (LAMB 1977) 99. Stercocaulan glahrum (Mül!. Arg.) Vain. (LINDSAY 1975, LAMB 1977, HUNECK er al. 1986, 10GENSEN 1986) 100. Usnca (Neuropogon} acromelana Stirt. (LA MB 1964, WALKER 1985) 101. Usnea (NewopogoJl) antorctica DR. [= USI/ea crassa Zammuto in Dodge, = Usnea crombiei Dodgc, = Usnea jloriformis Dodge, = Usnea granutifera (Vain.) Moryka, =Usnca propogullfera Dodgc, = Usnea psendofntticosa Zammuto in Dodge, = Usnea pustulata Dodge,? = Usnea subjoveolata Dodge,? =Usnea subpapillata Dodge] (CAMB 1964, GOLUBKOVA and SAVICZ 1970, SMITH 1972. WALKER 1985. HUNECK ct al. 1986) 102. Usnea tNcuropogon} aurantiaca-atra} (Jacq.) BOf)' [= Usnea auranüaca Motyka. = Usnea fascicüa Torrey, = Usnea melaxantha Ach, ? = Usnea sip!eiZammuto in Dodge] (LAMB 1964. SMITH 1972. WALKER 1985) 103. Usnea (Ncuropogon} subantarctica F.], Walker (WALKER 1985) 104. Vermcariae/acop/aca Vain. (SMITH 1972, 0VSTEDAL 1986b) 105. Vermcaria tesseIatu/a Nyl. (LAMB 1948a, SM1TH 1972, 0VSTEDAL 1986b) 106. ZahlhrucknereIIa patagonica Henssen (HENSSEN 1977)

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