Lichenologi~t 27(6): THE 463-471 LICHEN Ulrik (1995) GENUS CALOPLACA REGIONS SQ)CHTING* and Maria IN POLAR OLECH$ Abstract: Extensive material of Calopluca from Arctic and Antarctic regions has been critically examined. A list of 49 species is presented for Arctic regions. They are presumed to have a more or less circurnpolar distribution. Twenty-two species are listed from the Antarctic region, but about ten more, probably undescribed species, are present there. Abour one-third of the species in the Antarctic region are bipolar or widespread in cold regions; these include mainly terricolous and muscicolous species and none of them are maritime. It is assumed that migration of the bipolar or cosmopolitan species has taken place along the Andean mountain chain, whereas the maritime polar species have evolved separately in the two hemispheres. The Culoplaca species of the Antarctic region are provisionally assigned to the following distribution types: continental Antarctic, western Antarctic, insul-Antarctic and sub-Antarctic. Culoplucu exrecucu, C. suxicolu and C. phaeocarpellu are recorded as new to the Antarctic region. Culoplucu johnstonii (Dodge) Sochting & Olech, comb. nav., is established as the correct name of C. tenuis Ovstedal. Q 1995 The British Lichen Society Introduction Arctic and alpine regions are known to have a large number of Caloplaca species. However, there are few comprehensive treatments covering such regions (Hansen et al. 1987; Poelt & Hinteregger 1993). Phytogeographical studies are hampered by the sparse collecting of the more subtle species and the difficulties in identifying the species due to lack of suitable floras. In recent years more information has accumulated due to focused collecting in Greenland, Svalbard, Siberia and northern Fennoscandia. Although many taxonomic problems are still unsolved and a number of species are still undescribed it is now possible to get a broad overview of the distribution of a considerable number of Caloplaca species. Recent papers on the Caloplaca flora of the Antarctic region (see below) and our own continuing studies of extensive collections from Antarctica and the sub-Antarctic islands have made it possible to compare the polar floras of the two hemispheres. The present paper is based on published data and a number of as yet unpublished records from polar regions, which will eventually be treated more thoroughly. Phytogeographical compilations should be made with extreme care, as published data are severely burdened by misidentifications. Most data here are accordingly based on material we have studied ourselves. *Department DK-1353 SBotanical of Mycology, Botanical Institute, University of Copenhagen, 0. Farimagsgade Copenhagen K, Denmark. Institute, Jagellonian University, Ul. Lubicz 46, F-3 1-5 12 Krakow, Poland. 0024-2829/95/060463+09 $12.00/O 0 1995 The British Lichen 2D, Society 464 THE Vol. 27 LICHENOLOGIST Delimitation of the Polar Floras Arctic and sub-Arctic regions are found in Alaska and Canada, Greenland, Svalbard, northern Fennoscandia, European Russia, northern Siberia and the Russian far North-East. We have not attempted to draw a southern border for these regions, as the ranges of the species are still insufficiently known. Instead, the floras of Greenland and Svalbard, both well studied by us, have been used to define the northern polar element. Similarly, the southern polar element (hereafter referred to as Antarctic) is defined as the species occurring on the Antarctic continent and on the South Shetland Islands, areas that have recently been intensively studied by us. Species unknown from those areas, but recorded from other peri-Antarctic islands are not included. The Arctic Caloplaca Flora The Greenland Caloplaca flora has been thoroughly studied by Hansen et al. (1987). The data in Table 1 are based on this source, but with some modifications based on later studies (unpublished). The records of Caloplaca species from Svalbard are based on Sochting (1989 and unpublished records) and Elvebakk & Hertel (1995). The lichen flora of northern Fennoscandia is documented in Santesson (1993) and the species included here are those occurring in the provinces of Firmmark, Troms and Tome Lappmark. Table 1 also includes unpublished collections made by Sachting. At least 13 species not known from Greenland or Svalbard occur in those provinces, but are not included in Table 1. Records from North America are derived from the North American checklist (Egan 1987) and only in a few cases have been verified or changed by us. Accordingly, the occurrences in North America must be considered provisional. Records from the northernmost parts of the former Soviet Union were recently published by Sechting et al. (1992). Older records from those regions have largely been omitted if specimens have not been checked. The Antarctic Caloplaca Flora The data on the Antarctic species as presented in Table 1 are based on specimens studied by ourselves, as the literature on Antarctic lichens is often unreliable. Several of the records presented here have not previously been published, and we are presently studying about 10 more, probably undescribed, species from the Antarctic area. In the present paper we also include data from a number of islands in the sub-Antarctic that have a very similar flora. We are, however, aware that these islands can support additional species with a more northern distribution. Discussion of Polar Distributions Even though some areas of the Arctic, for northern Asia are insufficiently studied, it Caloplaca flora is more or less circumpolar. of saxicolous species, is highly influenced example North America and appears that the high Arctic The distribution, particularly by the occurrence of suitable 1995 Polar Caloplaca-Gchting & Olech 465 substrata. Muscicolous and terricolous species seem to be more uniformly distributed. The proximity of large, only slightly separated, continents is believed to cause the rather uniform flora, and the richness in species has been attributed to the ability of the flora to migrate southwards and survive there during the late Cenozoic glaciations. Sub-Arctic species may be more restricted in their range, but our knowledge about their distribution is so far insufficient for final conclusions. Twenty-one species are listed from the Antarctic region and with our present knowledge we estimate that about 10 more undescribed and possibly endemic taxa are present. About one-third of the listed Antarctic species also occur in the Arctic, consisting mainly of species growing on detritus and mosses, namely: C. ammiospila, C. citrina, C. phaeocarpella, C. tetraspora and C. tiroliensis. The species in this group are also recorded from the Central European Alps, apart from C. phaeocarpella, which may have been overlooked. They probably represent an Arctic-Antarctic-Alpine element, which migrated along the Andean mountain range, or the Malaysian-Papuan link (Lindsay 1977). Long-distance dispersal from the Arctic is another possibility, which, however, seems less likely (see below). Other non-maritime lichens in the more mesic parts of the Antarctic region, such as Cladonia species, have been shown to have a mostly wide distribution in cool regions (Stenroos 1993). This pattern has been attributed to longrange dispersal, which was, however, facilitated by occurrence in other continents of the Southern Hemisphere. Surprisingly the most common species on detritus in the Arctic, C. cerina, has so far not been found in the Antarctic region. There is presently no explanation for this. Further studies on alpine Caloplaca in the tropics would serve to clarify the interpretation of the ‘bipolar’ distributions and possibly show this element to be cosmopolitan. None of the maritime species of the Northern Hemisphere have been found in the Antarctic region, and the majority of Caloplaca species endemic to the region are more or less maritime. It appears that maritime species have had little opportunity to migrate across the tropics and subtropics and must thus have evolved separately in the specific polar regions. The ‘endemism’ of polar maritime Caloplaca species is so striking that it can hardly be attributed to differences in the ecological conditions of the polar sea shores. Accordingly it appears that viable diaspores of the maritime species have not been capable of long-range dispersal between the two polar regions. We believe that there are historical reasons for the apparently isolated evolution of the floras of the two hemispheres. The antiquity of many Southern Hemisphere species was pointed out by KHmefelt (1990) based on distribution patterns of more northern species. Further speculation on the origin of the maritime Antarctic Caloplaca species must await critical study of related species on the Southern Hemisphere continents. Distribution Patterns in the Antarctic Region The Caloplaca species in the Antarctic region can be grouped provisionally according to their distribution, which reflects their ecological requirements as well as probably their history of dispersal. THE 466 TABLE LICHENOLOGIST 1. Caloplaca Vol. 27 species occurring in polar regions Polar Caloplaca species Arctic species alcaruln Poelt approxirnata (Lynge) H. Magn. arenaria (Pers.) Miill. Arg. borealis (Vain.) Poelt caesiorufella (Nyl.) Zahlbr. cascellana (Rashen) Poelt celata Th. Fr. cerina (Ehrh.) Th. Fr. concilians (Nyl.) H. Olivier decipens (Am.) Blomb. 81 Forss. diphyodes (Nyl.) Jatta epiphyca Lynge epithallina Lynge Javovirescens (Wulf.) Dalla Terre & Samth. fraudans (Th. Fr.) Oliv. jidvolutea (Nyl.) Jatta insularis Poelt invadens Lynge jemclandica H. Magn. jungemzanniae (Vahl) Th. Fr. lacrea (Massal.) Zahlbr. leptocheila H. Magn. lithophila H. Magn.* livida (Hepp) Jatta tnagni-jlii Poelt nivalis (Koerb.) Th. Fr. noeisii Sechting ad interim paulii Poelt psorocida Hansen, Poelt & Sechting pyracea (Ach.) Th. Fr.4 saxifrogonon Poelt scopularis (Nyl.) Lenau scoroplaca (Nyl.) H. Magn. sibirica H. Magn. sinapisperma (Lam. & DC.) Maheu & Gillet soropelta (Hansen, Poelt & Sochting) Sechting spicsbergensis H. Magn. zonlinii Savicz tomoksis H. Magn. rrachyphylla (Tuck.) Zahlbr. verruculifera (Vain.) Zahlbr. ‘Bipolar’ species annniospila (Ach.) Oliv. anchon-phoeniceon Poelt & Clauzade cinina (Hoffm.) Th. Fr.§ exsecuta (Nyl.) Dalla Tome & Sam&t locations* Gr Sv NA Si ELI + + + + + + + -I+ + + + + + + + + + -I+ + + + + + + + + + + f + -I- + + + + + + + + + + + + + + + + + + + + -I- + + + + + + + + + + + + + + -I+ + + + + + + + + + + + + + + + + + + An + + + + + + + + + + + + + + + + + + + + -I- + + + + + + -I- + + + + + + + + + + + + + + + + -I- + + + + + -I+ + + + ? + + + + + + + + + Continued + + + + 1995 Polar Caloplaca-Smhting TABLE 467 & Olech 1. Continued Polar locations* species Gr Sv NA Si Eu An ‘Bipolar’ species continued phaeocarpella (Nyl.) Zah1br.t saxicola (Hoffm.) Nordint tetraspora (Nyl.) Oliv. riroliensis Zahlbr. + + + + + + + + ? + + + + ? + + + + + + + + + + Caloplaca Antarctic species arhallina Darb. austrosheclandica (Zahlbr.) Olech & Sochting buelliae Olech & Sochting cirrochrooides (Vain.) Zahlbr. coralhgera (Hue) Zahlbr. iomma Olech & Sechting isidioclada Zahlbr. johnstonii (Dodge) Sochting & Olechf lucens (Nyl.) Zahlbr. millegrana (Mull. Arg.) Zahlbr. psoromatis Olech & Sochting regalis (Vain.) Zahlbr. siphonospora Olech L? Sochting sublobulata (Nyl.) Zahlbr. + + + + + + + + + + + + + + *Gr: Greenland; Sv: Svalbard; NA; North America; Si: Siberia; Eu: Europe; An: Antarctic region. *Species belonging to the C. holocarpa group are here separated into saxicolous forms, C. lirhophila and corticolous and lignicolous forms, C. Dracea. This separation must be regarded as highly provisional. Specimens of the C. holocarpa group growing in the Antarctic region probably belong to an undescribed taxon. §Caloplaca cirrina is variable and badly understood. Antarctic material may belong to several taxa, probably including the Northern Hemisphere taxon C. cimna. TCaloplaca johnsronii (Dodge) Sochting & Olech, comb. nav. Blastenia johnsronii Dodge, British, Australian, New Zealand Anrarcric Research Expedition 1929-31. 7: 223 (1948). Type: Crozet Islands, Possession Island, American Bay. 3 November 1929, BZO-19 (FH)-Syn.: Caloplaca tenuis 0vsteda1, Norsk Polarinzitutt .Skr$cer 185: 44 (1985). +&Species new to the Antarctic region. 1. Continental Antarctic Continental Antarctic species are widespread on all sides of the continent where suitable substrata are available, but are present also on the Antarctic Peninsula and on the South Shetland Islands (Fig. 1A). This group includes C. athallina and C. citrina, two species confined to often eutrophicated cushions of mosses. Caloplaca citrina in the Antarctic region is poorly understood and it is possible that it includes several taxa, at least on the Antarctic islands. 2. Western Antarctic Western Antarctic speciesare absent from the continent. They are restricted to the Antarctic Peninsula, South Shetland Islands, South Orkney Islands, THE 468 Vol. 27 LICHENOLOGIST Tristan b”lh ocoas Prince CamphIl 7 . Tristan 1.. Da Cunho Aucklnnd I. LJl&a// Edward - South \ Ds w Cunho Sandwich 1. Prince I. Edwnrd 1. Tasm=i” . D Trisrnn Dn Cunhn AftiC” outh Georgia A- ..So”lh : Snndwich I. Prince Edwnrd I alkl a South ‘.. SAt 1. Shcllond 1. sit, I.‘.. Aucklnnd FIG. 1 Distribution of four species A, C. athallina (continental Antarctic); (insuL4nrarctic); of Calopluca representing different B, C. regalis (western Antarctic); D, C. sublobulacu (SubAntarctic). 1. ‘bma”is distribution types. C, C. cirrochrooides Tierra de1 Fuego and Patagonia (Fig. 1B). Caloplaca regalis, representing this group, is a very conspicuous and much collected lichen. Accordingly, its range is very reliably known. Calopluca austroshetlundica is only recorded from the Peninsula, South Shetland Islands and South America. The western Antarctic area is considered to have been ice-free for the longest period of time (Lindsay Polar Caloplaca-Stichting 1995 & Olech 469 1977) and possibly the location where evolution of the species occurred. The islands further east of South America may have a similar climate, but have a shorter history and may not yet have been colonized by all the species with a potential ability to grow there. Furthermore, there may be fewer suitable niches due to their smaller size (Engelsksjon & Jorgensen 1986). Caloplaca regalis is largely found on shores fertilized by penguins, whereas C. austroshetlandica is a pioneer species on rocks in late melting sites. 3. Insul-Antarctic Insul-Antarctic species occur in the oceanic climate of the sub-Antarctic islands and have spread far to the East, even to Kerguelen. However, they have not yet been found in South America, South Africa or Australia (Fig. 1C). It may be significant that these species are spreading probably mainly by rather large diaspores, such as soredia, isidia and isidia-like thallus branches. Similar observations were made when comparing macrolichens from the South Orkney Islands with those that had reached the South Sandwich Islands (Lindsay 1977). Caloplaca cirrochrooides is a typical representative of this group, but a similar pattern is shown by C. isidioclada, which is found as far North as Tristan da Cunha (37”15’S, 12”13’E) and Juan Femindez Island (345, 78-81”W). Caloplaca coralligera, also belonging to this group, has been recorded as far south as Ross Island (785). Caloplaca cirrochrooides and C. coralligera are maritime species, and all three species grow mainly on heavily fertilized rocks. 4. Sub-Antarctic Sub-Antarctic species are found on the sub-Antarctic islands, but do not occur on the Antarctic continent or on the Antarctic peninsula. They have a more northern distribution ranging even to the southern shores of South America, Africa, Australia or New Zealand (Fig. 1 D) . The northern limits of distribution of these species are in need of investigation. Most widespread is C. sublobulata, reaching all the three continents. Caloplaca hens reaches South America and C. millegrana is recorded from Australia. All three species are maritime, growing mainly on eutrophicated rocks. 5. Endemic Endemic species are so far known from the South Shetland Islands, namely C. buelliae, C. iomma, C. psoromatis and C. siphonospora. They may subsequently be found in other parts of Antarctica, so it is premature to discuss their distribution. Discussion of Antarctic Distribution Patterns Phytogeographical zones of the Southern Hemisphere were discussed by Galloway (1991). He recognized an Antarctic realm including an insulAntarctic subrealm with a very oceanic climate and a continental Antarctic subrealm consisting of the Antarctic continent with very arid climatic 470 THE LICHENOLOGIST Vol. 27 conditions. The original circumscription of the insul-Antarctic sub-realm (Fleming 1987) excluded the South Shetland Islands and the Antarctic Peninsula, two regions with an intermediate position which, at least according to the Galoplaca flora, do have strong affinities to the sub-Antarctic islands included in &he insul-Antarctic subrealm. Hertel (1984) studied the saxicdlous lecideoid lichens of the sub-Antarctic. He concluded provisionally that the continental flora consists of rather widespread species, whereas the subAntarctic islands have a more specific flora. The present study indicates that species not confined to maritime habitats are more widespread. Species associated with maritime climates, particularly on the sub-Antarctic islands, are largely specific to the Antarctic area and have probably evolved there. It may be speculated that they have survived on some of the western islands during the late glaciations and subsequently spread towards the East onto more recently developed islands. Such a process is supported by the studies of lecideoid lichens on Prince Edward Islands (Hertel 1984). Engelsksjan & Jrargensen (1986) emphasized the importance of the strong westerly winds that are able to transport diaspores from the western Antarctic region towards the more eastern and younger islands. Sea currents in the region have the same direction and may also be able to transport diaspores. We acknowledge loan of material from the following herbaria: FH, H, S and UPS, and wish to thank Professor H. Hertel and Dr C. Wetrnore for letting us study their collections. Dr R. Seppelt revised the language. He and Professor P. M. Jorgensen are thanked for valuable commems on the manuscript. Ruth Bruus Jakobsen and Else Meier Andersen prepared the maps. REFERENCES Elvebakk, A. & Hertel, H. (1995) Part VI. Lichens. In A Cafalogue ofrhe Svalbard Plants, Fungi, Algae, and Cyanobacteria (A. Elvebakk & P. Prestmd, eds). Norsk Polarinstitutr SknjIer 198 (in press). Engelsksjen, T. &Jsrgensen, P. M. (1986) Phytogeographical relations of the cryptogamic flora of Bouveteya. Norsk Polarinstimz Sknj?eer 185: 71-79. Egan, R. S. (1987) A fZ.h checklist of the lichen-forming, lichenicolous and allied fungi of the continental United Stated and Canada. Byologist 90: 77-173. Fleming (1987) Comments on Udvardy’s biogeographical realm Antarctica. 3ournal of the Royal Society of New Zealand 17: 195-200. Galloway, D. J. (1991) Phytogeography of southern hemisphere lichens. In Quantitative Approaches IO Phyrogeography (P. L. Nirnis & T. J. Crovello, eds): 233-262. Dordrecht: Kluwer. Hansen, E. S., Poelt, J. 8c Sochting, U. (1987) Die Flechtengattung Caloplaca in GrBnland. Meddr Grmland, Bioscience 25: l-52. Herrel, H. (1984) iiber saxicole, lecideoide Flechten der Subanrarktis. Beihefr Nova Hedwiflb 79: 399-499. K;imefelt, I. (1990) Evidence of a slow evolutionary change in the speciation of lichens. Bibliotheca Lichenologiia 38: 291-306. Lindsay, D. C. (1977) Lichens of cold deserts. In Lichen Ecology (M. R. D. Seaward, ed.): 183-209. London: Academic Press. Olech, M. & Sachring, U. (1993) Four new species of Caloplaca from Antarctica. Lichenologisr 25: 261-269. Poelt, J. & Hinteregger, E. (1993) Beitrege zur Kennmis der Flechtenflora des Himalaya. VII Die Gattungen Caloplaca, Fulgensib und Ioplaca. Bibliotheca Lichenologiia 50: l-247. Santesson (1993) The lichens and lichenicolousfungi of Sweden and Norway. Lund: SBT-fdrlaget. Steenroos, S. (1993) Taxonomy and distribution of the lichen family Cladoniaceae in the Antarctic and peri-Antarctic regions. Cryproganlic Botany 3: X0-344. 1995 Polar Caloplaca-Smhting 6, Olech 471 Sachting (1989) Lignicolous species of the Lichen genus Calopluca from Svalbard. Opera Boranica 100: 241-257. Sechting, U. 81 0vsteda1, D. 0. (1992) Contributions to the Calopluca flora of the western Antarctic region. Nordic Journal Botany 12: 121-134. Scxhting, U., Zhurbenko, M. & Hansen, E. S. (1992) Notes on the genus Caloplaca in the Siberian arctic. Graphis Scrtpta 4: 30-32. Accepted for publication 24 May 1995