Author s personal copy. Terricolous Lichens in the Glacier Forefield of the Pasterze (Eastern Alps, Carinthia, Austria)

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3 Phyton (Horn, Austria) Vol. 55 Fasc DOI: /0380.phyton55(2) Terricolous Lichens in the Glacier Forefield of the Pasterze (Eastern Alps, Carinthia, Austria) By Peter O. Bilovitz*), Anja Wallner, Veronika Tutzer, Juri Nascimbene**) and Helmut Mayrhofer*) With 1 Figure Received April 30, 2015 Key words: Lichenized Ascomycetes, Lichenes. Biodiversity, ecology, flora, floristics. Alps, glacier forefield, glacier retreat. Summary Bilovitz P. O., Wallner A., Tutzer V., Nascimbene J. & Mayrhofer H Terricolous lichens in the glacier forefield of the Pasterze (Eastern Alps, Carinthia, Austria). Phyton (Horn, Austria) 55 (2): , with 1 figure. The investigation of lichens on soil, plant debris and terricolous mosses in the glacier forefield of the Pasterze yielded 35 lichen species. Placidiopsis oreades Breuss (Verrucariales) is new to Austria. Three sampling sites were established at increasing distance from the glacier, in order to compare species diversity, abundance and composition within the forefield and with four other glacier forefields of the Eastern Alps. Zusammenfassung Bilovitz P. O., Wallner A., Tutzer V., Nascimbene J. & Mayrhofer H Terricolous lichens in the glacier forefield of the Pasterze (Eastern Alps, Carinthia, Austria). [Terricole Flechten im Gletschervorfeld der Pasterze (Ostalpen, Kärnten, Österreich)]. Phyton (Horn, Austria) 55 (2): , mit 1 Abbildung. *) Mag. Dr. Peter O. Bilovitz (corresponding author), Ao. Univ.-Prof. Mag. Dr. Helmut Mayrhofer, Institute of Plant Sciences, NAWI Graz, University of Graz, Holteigasse 6, 8010 Graz, Austria, Europe; pe.bilovitz@uni-graz.at, helmut. mayrhofer@uni-graz.at **) Dr. Juri Nascimbene, Department of Agronomy, Food, Natural Resources, Animals and the Environment (DAFNAE), University of Padova, viale dell Università 16, 35020, Legnaro, Padova, Italy, Europe; juri.nascimbene@unipd.it

4 202 Die Erhebung der Flechten auf Erde, Pflanzenresten und terricolen Moosen im Gletschervorfeld der Pasterze ergab 35 Flechtenarten. Placidiopsis oreades Breuss (Verrucariales) ist ein Neufund für Österreich. Es wurden drei Aufnahmeflächen mit zunehmender Entfernung vom Gletscher festgelegt, um Diversität, Abundanz und Zusammensetzung der Arten innerhalb des Vorfeldes und mit vier weiteren Gletschervorfeldern der Ostalpen zu vergleichen. 1. Introduction Glacier retreat on a global scale started with the end of the Little Ice Age around Since 1980, a significant global warming has led to glacier retreat becoming increasingly rapid and ubiquitous. In the past decades, the majority of glaciers in the Alps has experienced considerable mass losses. Currently, around 0.5 % of the Austrian territory is covered by ice (Sulzer & Lieb 2009). In the glacier year 2012/2013, 90 % of the surveyed Austrian glaciers retreated (Fischer 2014), also the Pasterze, with an average length loss of 41 metres (2012: 97.3 m; 2011: 40.3 m). The growing areas of recently bared glacier forefields are providing new ecological niches for pioneer organisms such as soil lichens, but little is known about their colonization patterns in these special habitats. In the framework of a project on the impact of changing local conditions on lichen occurrence in glacier retreat regions, we investigated the terricolous lichen biota of five glacier forefields in the Eastern Alps (see also Bilovitz & al. 2014a, 2014b, 2014c, 2015). The floristic data from the forefield of the Pasterze glacier (Fig. 1) in Carinthia, Austria are presented in this paper. 2. Investigation Area With a current length of about 8 km, the Pasterze is the longest glacier in the Eastern Alps, situated within the Hohe Tauern mountain range in the province of Carinthia, directly beneath Austria s highest mountain, the pyramid-shaped Großglockner (3798 m). The Pasterze can be easily reached via the Großglockner High Alpine Road, which connects the provinces Salzburg and Carinthia. The Kaiser-Franz-Josefs-Höhe visitors centre at 2369 m offers a panoramic view over the Pasterze with its forefield and the surrounding mountains. The glacier forefield of the Pasterze is situated within the so-called Tauern window, a geological structure, where high-grade metamorphic rocks of the underlying Penninic nappes crop out. The dominant rock types around the Pasterze mainly comprise calcareous mica-schists and greenschists (= prasinite). Detailed information about the geology of the Glockner Group can be found in Frank 1969, Krainer 1988 and Krainer Burger & Franz 1969 gave an overview on the soil development of the Pasterze area. The Glockner Group is situated in the transition zone of the atlanticcontinental climate regime (Tollner 1969). According to Paschinger 1976,

5 203 Fig. 1. Pasterze glacier and its forefield, Johannisberg in the background. Phot. P. O. Bilovitz, 05.IX.2013.

6 204 three climate types can be distinguished: the continental valley climate, the weak continental slope climate and the atlantic frost climate of higher altitudes. According to this, the Tauern valleys are relatively dry, for example Heiligenblut (1380 m) with an average annual precipitation of 825 mm ( , Tschernutter 1982), whereas the continental character of the climate becomes weaker with increasing elevation. Standard measurements of the Pasterze glacier have been conducted annually since 1879, thus the amount of data is remarkable. Since the end of the Little Ice Age, the glacier has retreated almost continuously, only interrupted by few periods of stagnation or slight advances (Sulzer & Lieb 2009). The discovery of parts of coniferous trees and pieces of peat (dated back mainly to the early Holocene) in the area of the Pasterze forefield, washed out by the stream from under the present ice, provided indications of smaller stages of the glacier in postglacial time (Slupetzky 1993, Slupetzky & al. 1998, Nicolussi & Patzelt 2000, Nicolussi & Patzelt 2001, Drescher-Schneider & Kellerer-Pirklbauer 2008). The Hohe Tauern mountain range, in particular the area of the Großglockner and the surroundings of the Pasterze, has been of special interest for botanists for a long time. Details on the natural history exploration of the Glockner Group were provided by Gams 1936 in the first part of his paper. The second part deals with the living conditions, the third part with the flora and the fourth part with the vegetation of the area, followed by an overview of the plant communities plus a vegetation map. Friedel 1956 published a vegetation map of the environs of the Pasterze together with a comprehensive annotation. The map represents the state of the area in the year At that time, the glacier reached the Elisabethfelsen (2156 m). Friedel 1969 presented a short contribution on the vegetation of the area with an enclosed vegetation map. A more recent vegetation map of the Großglockner area is provided by Schiechtl & Stern Böhm 1969 dealt with the timber line of the Glockner Group. A contribution on the vegetation development of the forefield of the Pasterze was presented by Zollitsch 1969, whereas Franz 1969 reported on the colonization of the forefield with invertebrates. The local flora and vegetation of the Pasterze glacier forefield was presented and discussed in respect to glaciology and vegetation dynamics by Wittmann & al. 2009, dealing with the following plant communities: Seslerio-Caricetum sempervirentis, Rhododendretum ferruginei, Salicetum retusae-reticulatae, Salicetum helveticae, Salicetum waldsteinianae, Dryadetum octopetalae, Drabion hoppeanae, Caricetum frigidae as well as two rare communities of alpine alluvial habitats, namely the Carex bicolor-community and the Carex atrofusca-community; additionally, the Astero bellidiastro-kobresietum simpliciusculae was outlined. A remarkable locality is the Gamsgrube, situated above the Pasterze within a southwest exposed massive bowl-shaped cirque of the Fuscherkarkopf. For the most part, it is covered with wind-driven sand, thus provid-

7 205 ing unique living conditions in the Alps (e. g. Friedel 1951, Gams 1951, Hartl 1988). The Gamsgrube is the locus classicus of Braya alpina Sternb. & Hoppe (Sternberg & Hoppe 1815), a rare and endangered endemic species of the Eastern Alps (Fischer & al. 2008). A compilation of the lichens of Carinthia was presented by Türk & al Türk & Hafellner 1992 provided a comprehensive list of 660 lichen species and Hafellner & Türk 1995 a list with more than 140 lichenicolous fungi both lichenized and non-lichenized, occurring in the Carinthian part of the Hohe Tauern National Park. Egger 1997 surveyed site dynamics and succession of lichen colonization in the glacier forefield of the Winkelkees in the Seebach valley, situated in the Carinthian part of the Hohe Tauern National Park. 3. Material and Methods Sampling location: Austria, Carinthia, Hohe Tauern, Glockner Group, Hohe Tauern National Park, NW of Heiligenblut, N/ E, m, glacier forefield of the Pasterze, 05.IX.2013, leg. P. Bilovitz, V. Tutzer & A. Wallner. Three sampling sites were established at increasing distance from the glacier, corresponding to a gradient of moraine age: site 1 = c. 600 m (ice-free for c. 30 years), site 2 = c m (ice-free for c. 35 years), site 3 = c m (ice-free for c. 60 years). In each site, lichens were surveyed within five 1 x 1 m randomly placed plots, both on soil and on plant debris or decaying terricolous mosses. Spots with larger stones were avoided. Phanerogams were present in all three sites, but, with increasing distance from the glacier, diversity rose and vegetation cover became significantly denser. Each plot was divided into 10 x 10 cm quadrats (Bilovitz & al. 2014a: Fig. 2), in order to obtain data on species frequency (max. frequency/plot = 100). For each species, specimens were collected for a more accurate identification in the laboratory. The specimens have been identified mainly with the aid of Wirth & al. 2013, using routine light microscopy techniques. Some of the identifications required verification by using standardized thin-layer chromatography (TLC), following the protocols of White & James 1985 and Orange & al The specimens are preserved in the herbarium of the Institute of Plant Sciences, University of Graz (GZU). The nomenclature mainly follows Wirth & al. 2013, or other modern treatments. 4. Results and Discussion In total, 35 lichen species were found in the three sampling sites (Table 1). A number of lichens showed the influence of calcareous substrate, for instance Allocetraria madreporiformis and Vulpicida juniperinus. The diversity of terricolous lichens near the front of the glacier was very low. The first lichens occurred at a distance of about 600 m from the front (site 1), and only 9 species were found in this sampling site. The foliose lichen Peltigera rufescens was the most frequent species (frequency of 69). The rest of the species only occurred with very low frequency values (1 3).

8 206 At a distance of about 1000 m to the glacier (site 2), the number of species increased significantly to 23. Fulgensia bracteata subsp. deformis reached the highest frequency value (62), followed by Caloplaca stillicidiorum s. l. (38), C. tiroliensis (37), Lecanora hagenii var. fallax (35), Cladonia pyxidata s. l. (35), Allocetraria madreporiformis (20), Stereocaulon alpinum (19), Peltigera rufescens (15) and Flavocetraria nivalis (6). The rest of the species only occurred with very low frequency values (1 3). At a distance of about 1300 m from the glacier (site 3), we observed a further increase in species diversity. In total, 32 species were found in this site. The fruticose lichens Cladonia pyxidata s. l. (122), Cladonia symphycarpia (87) and Stereocaulon alpinum (71) reached the highest frequency values, followed by Fulgensia bracteata subsp. deformis (61), Caloplaca tiroliensis (50), Lecanora hagenii var. fallax (47), Physconia muscigena (43), Caloplaca stillicidiorum s. l. (32), Peltigera rufescens (28), Vulpicida juniperinus (26), Cetraria muricata (20), Thamnolia vermicularis (16), Flavocetraria nivalis (15), Allocetraria madreporiformis (10) and Lecanora epibryon (10). The rest of the species occurred with frequency values less than ten. Table 1. List of lichenized taxa with their substrata and the frequency of each species in the three sampling sites (deb = on plant debris or decaying terricolous mosses; ter = on soil). Frequency Taxon Substratum Site 1 Site 2 Site 3 Allocetraria madreporiformis (Ach.) Kärnefelt & A. Thell ter Bacidia bagliettoana (A. Massal. & De Not.) deb Jatta Bilimbia lobulata (Sommerf.) Hafellner & Coppins Blennothallia crispa (Huds.) Otálora, P. M. Jørg. & Wedin ter ter Caloplaca stillicidiorum s. l. deb Caloplaca tiroliensis Zahlbr. deb Cetraria ericetorum Opiz ter Cetraria islandica (L.) Ach. ter Cetraria muricata (Ach.) Eckfeldt ter Cladonia macroceras (Delise) Hav. ter Cladonia pyxidata s. l. ter * Cladonia symphycarpia (Flörke) Fr. ter Diploschistes muscorum (Scop.) R. Sant. Cladonia sp

9 207 Taxon Substratum Frequency Site 1 Site 2 Site 3 Flavocetraria cucullata (Bellardi) Kärnefelt & A. Thell ter Flavocetraria nivalis (L.) Kärnefelt & A. Thell ter Fulgensia bracteata (Hoffm.) Räsänen subsp. deformis (Erichsen) Poelt ter Hypogymnia physodes (L.) Nyl. deb Lecanora bryopsora (Doppelb. & Poelt) Hafellner & Türk deb Lecanora epibryon (Ach.) Ach. deb Lecanora hagenii (Ach.) Ach. var. fallax Hepp deb Lecidella wulfenii (Hepp) Körb. deb Lepraria eburnea J. R. Laundon deb Megaspora verrucosa (Ach.) Hafellner &V. Wirth deb Peltigera rufescens (Weiss) Humb. ter Physconia muscigena (Ach.) Poelt ter Placidiopsis oreades Breuss ter Pseudevernia furfuracea (L.) Zopf var. ceratea (Ach.) D. Hawksw. Rinodina candidogrisea Hafellner, Muggia & Obermayer deb deb Rinodina roscida (Sommerf.) Arnold deb Scytinium intermedium (Arnold) Otálora, P. M. Jørg. & Wedin ter Solorina bispora Nyl. ter Stereocaulon alpinum Laurer ter Stereocaulon nanodes Tuck. ter Thamnolia vermicularis (Sw.) Schaer. ter Vulpicida juniperinus (L.) J.-E. Mattsson & M. J. Lai *) doubtful frequency data. ter In comparison to the forefields of the Morteratsch glacier in Graubünden (Bilovitz & al. 2015) with 13 lichen species, the Rötkees in South Tyrol (Bilovitz & al. 2014a) with 29 species, the Matscherferner in South Tyrol (Bilovitz & al. 2014b) with 34 species and the Gaisbergferner in Tyrol (Bilovitz & al. 2014c) with 39 species, the total diversity of the Pasterze forefield is similar

10 208 to that of the Matscherferner and the Gaisbergferner. Considering the species composition, the glacier forefield of the Gaisbergferner is most similar to that of the Pasterze, 21 out of 53 species being shared (Table 2). The number of lichen species in all five glacier forefields adds up to a total of 83. Five species occurred in each of them, namely Cetraria islandica, Cladonia macroceras, C. pyxidata s. l., Peltigera rufescens and Stereocaulon alpinum. The latter is a fruticose species and presumably the most constant soil lichen occurring in glacier forelands of the Alps (see Bilovitz & al. 2015). In contrast to the four other investigated glacier forefields in the Eastern Alps (Bilovitz & al. 2014a, 2014b, 2014c, 2015), Stereocaulon alpinum was not the most frequent species in the Pasterze forefield. The soil-inhabiting squamulose lichen Placidiopsis oreades Breuss (Verrucariales) is new to Austria. According to Breuss 1996, the species has a scattered distribution and is known from the Alps, the Carpathians and Kirghizia. In the Alps there are just few records, namely from Bavaria (Germany) and the canton Bern (Switzerland). The commonly sterile sorediate lichen Rinodina candidogrisea, a recently described species, seems to be relatively common in the Alps, where it occurs from the upper montane to the alpine vegetation belt (Hafellner & al. 2012). To date, the species is known from the Austrian provinces Carinthia, Styria, Upper Austria and Vorarlberg, the German state Bavaria, the Italian provinces Friuli-Venezia Giulia, Piemonte and Trentino-Alto Adige, Slovenia and the Swiss canton Graubünden. The results in this paper as well as the results in Bilovitz & al. 2014a, 2014b, 2014c, 2015 show that increasing lichen diversity and abundance directly correlate with the increasing age of the moraine. Similar findings were presented by Türk & Erschbamer 2010a, 2010b, who listed 31 lichens growing on soil, plant debris and terricolous mosses from the Rotmoosferner in Tyrol and found the same pattern of lichen diversity in relation to moraine age. Table 2. Terricolous lichens in the glacier forefield of the Morteratsch glacier (1; Bilovitz & al. 2015), Rötkees (2; Bilovitz & al. 2014a), Matscherferner (3; Bilovitz & al. 2014b), Pasterze (4; this paper), Gaisbergferner (5; Bilovitz & al. 2014c). Taxon Alectoria ochroleuca (Hoffm.) A. Massal. Allocetraria madreporiformis (Ach.) Kärnefelt & A. Thell Amandinea punctata (Hoffm.) Coppins & Scheid. Arthrorhaphis citrinella (Ach.) Poelt Arthrorhaphis spec. [A. alpina or A. vacillans] Bacidia bagliettoana (A. Massal. & De Not.) Jatta

11 209 Taxon Bilimbia lobulata (Sommerf.) Hafellner & Coppins Bilimbia microcarpa (Th. Fr.) Th. Fr. Blennothallia crispa (Huds.) Otálora, P. M. Jørg. & Wedin Bryonora castanea (Hepp) Poelt Caloplaca ammiospila (Wahlenb.) H. Olivier Caloplaca sinapisperma (Lam. & DC.) Maheu & Gillet Caloplaca stillicidiorum s. l. Caloplaca tiroliensis Zahlbr. Catapyrenium cinereum (Pers.) Körb. Cetraria ericetorum Opiz Cetraria islandica (L.) Ach. Cetraria muricata (Ach.) Eckfeldt Cladonia amaurocraea (Flörke) Schaer. Cladonia arbuscula s. l. Cladonia arbuscula (Wallr.) Flot. subsp. squarrosa (Wallr.) Ruoss Cladonia borealis S. Stenroos Cladonia cariosa s. l. Cladonia fimbriata (L.) Fr. Cladonia gracilis (L.) Willd. Cladonia macroceras (Delise) Hav. Cladonia cf. mitis Sandst. Cladonia cf. pleurota (Flörke) Schaer. Cladonia pyxidata s. l. Cladonia cf. subulata (L.) Weber ex F. H. Wigg. Cladonia symphycarpia (Flörke) Fr. Cladonia uncialis (L.) Weber ex F. H. Wigg. Cladonia spec. Dactylina ramulosa (Hook.) Tuck. Dibaeis baeomyces (L. f.) Rambold & Hertel Diploschistes muscorum (Scop.) R. Sant. Flavocetraria cucullata (Bellardi) Kärnefelt & A. Thell Flavocetraria nivalis (L.) Kärnefelt & A. Thell

12 210 Taxon Fulgensia bracteata (Hoffm.) Räsänen subsp. deformis (Erichsen) Poelt Fuscopannaria praetermissa (Nyl.) P. M. Jørg. Hypogymnia physodes (L.) Nyl. Lecanora bryopsora (Doppelb. & Poelt) Hafellner & Türk Lecanora epibryon (Ach.) Ach. Lecanora hagenii (Ach.) Ach. var. fallax Hepp Lecidea berengeriana (A. Massal.) Th. Fr. Lecidea hypnorum Lib. Lecidella wulfenii (Hepp) Körb. Lecidoma demissum (Rutstr.) Gotth. Schneid. & Hertel Lepraria diffusa (J. R. Laundon) Kukwa Lepraria eburnea J. R. Laundon Lepraria finkii (Hue) R. C. Harris Megaspora verrucosa (Ach.) Hafellner & V. Wirth Micarea incrassata Hedl. Ochrolechia inaequatula sensu auct. Peltigera didactyla (With.) J. R. Laundon Peltigera extenuata (Nyl. ex Vain.) Lojka Peltigera lepidophora (Nyl. ex Vain.) Bitter Peltigera rufescens (Weiss) Humb. Phaeorrhiza nimbosa (Fr.) H. Mayrhofer & Poelt Physconia muscigena (Ach.) Poelt Placidiopsis oreades Breuss Placynthiella icmalea (Ach.) Coppins & P. James Protopannaria pezizoides (Weber) P. M. Jørg. & S. Ekman Protothelenella sphinctrinoidella (Nyl.) H. Mayrhofer & Poelt Pseudevernia furfuracea (L.) Zopf var. ceratea (Ach.) D. Hawksw. Psoroma hypnorum (Vahl) Gray Psoroma tenue Henssen var. boreale Henssen Pycnothelia papillaria (Ehrh.) Dufour

13 211 Taxon Rinodina candidogrisea Hafellner, Muggia & Obermayer Rinodina mniaraea (Ach.) Körb. var. mniaraea Rinodina mniaraea (Ach.) Körb. var. cinnamomea Th. Fr. Rinodina mniaraea (Ach.) Körb. var. mniaraeiza (Nyl.) H. Magn. Rinodina roscida (Sommerf.) Arnold Scytinium intermedium (Arnold) Otálora, P. M. Jørg. & Wedin Solorina bispora Nyl. Solorina crocea (L.) Ach. Sporodictyon terrestre (Th. Fr.) S. Savić & Tibell Stereocaulon alpinum Laurer Stereocaulon nanodes Tuck. Tetramelas insignis (Nägeli) Kalb Thamnolia vermicularis (Sw.) Schaer. var. vermicularis Thamnolia vermicularis (Sw.) Schaer. var. subuliformis (Ehrh.) Schaer. Toninia spec. Trapeliopsis granulosa (Hoffm.) Lumbsch Vulpicida juniperinus (L.) J.-E. Mattsson & M. J. Lai sterile, sorediate crustose lichen 1 sterile, sorediate crustose lichen 2 5. Acknowledgements We would like to thank the administration of the National Park Hohe Tauern Carinthia and the Amt der Kärntner Landesregierung (Abteilung 8) for supporting our request for a scientific collecting permit, the Bezirkshauptmannschaft Spittal an der Drau (Bereich 10) for making out the permit, Peter Kosnik for the TLC, Othmar Breuss for the confirmation of Placidiopsis oreades, and Christian Scheuer as well as Herwig Teppner for critically reading the manuscript and general remarks. Financial support from the Austrian Science Fund (FWF project P25078-B16) and the University of Graz is gratefully acknowledged.

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