Author’s personal copy

Phyton (Horn, ) Vol. 55 Fasc. 2 201–214 15. 12. 2015

DOI: 10.12905/0380.phyton55(2)2015-0201

Terricolous Lichens in the Glacier Forefield of the Pasterze (, , 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. 2015. Ter- ricolous lichens in the glacier forefield of the Pasterze (Eastern Alps, Carinthia, Austria). – Phyton (Horn, Austria) 55 (2): 201–214, 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, abun- dance 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. 2015. Ter- ricolous 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): 201–214, 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; e-mail: [email protected], helmut. [email protected] **) 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; e-mail: [email protected] Author’s personal copy

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 1850. 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 condi- tions on lichen occurrence in glacier retreat regions, we investigated the ter- ricolous 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 (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 pyr- amid-shaped Großglockner (3798 m). The Pasterze can be easily reached via the Großglockner High Alpine Road, which connects the provinces and Carinthia. The Kaiser-Franz-Josefs-Höhe visitors’ centre at 2369 m of- fers a panoramic view over the Pasterze with its forefield and the surround- ing mountains. The glacier forefield of the Pasterze is situated within the so-called Tau- ern 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 can be found in Frank 1969, Krainer 1988 and Krainer 1994. 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 atlantic- continental climate regime (Tollner 1969). According to Paschinger 1976, Author’s personal copy

203 , 05.IX.2013. , ilovitz B Fig. 1. Pasterze glacier and its forefield, Johannisberg in the background. – Phot. P. O. O. P. – Phot. in the background. Johannisberg glacier and its forefield, Pasterze 1. Fig. Author’s personal copy

204 three climate types can be distinguished: the continental valley climate, the weak continental slope climate and the atlantic frost climate of higher alti- tudes. According to this, the Tauern valleys are relatively dry, for example Heiligenblut (1380 m) with an average annual precipitation of 825 mm (1971–1980, 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 inter- rupted 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 small- er stages of the glacier in postglacial time (Slupetzky 1993, Slupetzky & al. 1998, Nicolussi & Patzelt 2000, Nicolussi & Patzelt 2001, Drescher-Schnei- der & Kellerer-Pirklbauer 2008). The Hohe Tauern mountain range, in particular the area of the Groß- glockner 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 pa- per. 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 byan overview of the plant communities plus a vegetation map. Friedel 1956 pub- lished a vegetation map of the environs of the Pasterze together with a com- prehensive annotation. The map represents the state of the area in the year 1934. 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 1985. Böhm 1969 dealt with the tim- ber line of the Glockner Group. A contribution on the vegetation develop- ment of the forefield of the Pasterze was presented by Zollitsch 1969, whereas Franz 1969 reported on the colonization of the forefield with inver- tebrates. 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: Sesler- io-Caricetum sempervirentis, Rhododendretum ferruginei, Salicetum re- tusae-reticulatae, Salicetum helveticae, Salicetum waldsteinianae, Dryade- tum octopetalae, Drabion hoppeanae, Caricetum frigidae as well as two rare communities of alpine alluvial habitats, namely the Carex bicolor-commu- nity and the Carex atrofusca-community; additionally, the Astero bellidias- tro-Kobresietum simpliciusculae was outlined. A remarkable locality is the Gamsgrube, situated above the Pasterze within a southwest exposed massive bowl-shaped cirque of the Fuscher­ karkopf. For the most part, it is covered with wind-driven sand, thus provid- Author’s personal copy

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. 2004. 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 Na- tional Park.

3. Material and Methods

Sampling location: Austria, Carinthia, Hohe Tauern, Glockner Group, Hohe Tauern National Park, NW of Heiligenblut, 47°04’N/12°44’–45’E, 2075–2090 m, gla- cier forefield of the Pasterze, 05.IX.2013, leg. P.B ilovitz, 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. 1000 m (ice-free for c. 35 years), site 3 = c. 1300 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 dis- tance 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 labora- tory. 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. 2001. The specimens are pre- served 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 (Ta- ble 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). Author’s personal copy

206

At a distance of about 1000 m to the glacier (site 2), the number of spe- cies increased significantly to 23. Fulgensia bracteata subsp. deformis reached the highest frequency value (62), followed by Caloplaca stillicidio- rum 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 symphy- carpia (87) and Stereocaulon alpinum (71) reached the highest frequency values, followed by Fulgensia bracteata subsp. deformis (61), Caloplaca tiro- liensis (50), Lecanora hagenii var. fallax (47), Physconia muscigena (43), Ca- loplaca stillicidiorum s. l. (32), Peltigera rufescens (28), Vulpicida juniperi- nus (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 spe- cies in the three sampling sites (deb = on plant debris or decaying terricolous moss- es; ter = on soil). Frequency Substra- Taxon Site 1 Site 2 Site 3 tum

Allocetraria madreporiformis (Ach.) Kärnefelt ter 0 20 10 & A. Thell

Bacidia bagliettoana (A. Massal. & De Not.) deb 0 1 1 Jatta

Bilimbia lobulata (Sommerf.) Hafellner & ter 0 0 3 Coppins

Blennothallia crispa (Huds.) Otálora, P. M. ter 3 1 0 Jørg. & Wedin Caloplaca stillicidiorum s. l. deb 1 38 32

Caloplaca tiroliensis Zahlbr. deb 0 37 50

Cetraria ericetorum Opiz ter 0 0 4

Cetraria islandica (L.) Ach. ter 2 2 1

Cetraria muricata (Ach.) Eckfeldt ter 0 0 20

Cladonia macroceras (Delise) Hav. ter 0 0 2 Cladonia pyxidata s. l. ter 0 35 122*

Cladonia symphycarpia (Flörke) Fr. ter 1 0 87

Diploschistes muscorum (Scop.) R. Sant. Cladonia sp. 0 0 5 Author’s personal copy

207

Frequency Substra- Taxon Site 1 Site 2 Site 3 tum

Flavocetraria cucullata (Bellardi) Kärnefelt ter 0 1 1 & A. Thell

Flavocetraria nivalis (L.) Kärnefelt & A. Thell ter 0 6 15

Fulgensia bracteata (Hoffm.) Räsänen subsp. ter 1 62 61 deformis (Erichsen) Poelt

Hypogymnia physodes (L.) Nyl. deb 0 2 1

Lecanora bryopsora (Doppelb. & Poelt) deb 0 0 9 Hafellner & Türk

Lecanora epibryon (Ach.) Ach. deb 0 1 10

Lecanora hagenii (Ach.) Ach. var. fallax Hepp deb 0 35 47

Lecidella wulfenii (Hepp) Körb. deb 0 0 1

Lepraria eburnea J. R. Laundon deb 0 0 7

Megaspora verrucosa (Ach.) Hafellner & V. Wirth deb 1 1 3

Peltigera rufescens (Weiss) Humb. ter 69 15 28

Physconia muscigena (Ach.) Poelt ter 0 2 43

Placidiopsis oreades Breuss ter 0 0 1

Pseudevernia furfuracea (L.) Zopf var. ceratea deb 0 1 1 (Ach.) D. Hawksw.

Rinodina candidogrisea Hafellner, Muggia & deb 0 2 1 Obermayer

Rinodina roscida (Sommerf.) Arnold deb 0 0 1

Scytinium intermedium (Arnold) Otálora, P. ter 0 2 0 M. Jørg. & Wedin

Solorina bispora Nyl. ter 0 3 1

Stereocaulon alpinum Laurer ter 1 19 71

Stereocaulon nanodes Tuck. ter 0 1 0

Thamnolia vermicularis (Sw.) Schaer. ter 1 0 16

Vulpicida juniperinus (L.) J.-E. Mattsson & ter 0 2 26 M. J. Lai

*) doubtful frequency data.

In comparison to the forefields of the in Graubünden (Bilovitz & al. 2015) with 13 lichen species, the Rötkees in South (Bilo- vitz & 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 Author’s personal copy

208 to that of the Matscherferner and the Gaisbergferner. Considering the spe- cies 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 (Ger- many) and the canton Bern (Switzerland). The commonly sterile sorediate lichen Rinodina candidogrisea, a re- cently 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 Carin- thia, 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 di- rectly 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 1 2 3 4 5

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 + + + Author’s personal copy

209

Taxon 1 2 3 4 5

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. squar- + + rosa (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 + + + Author’s personal copy

210

Taxon 1 2 3 4 5

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 + Author’s personal copy

211

Taxon 1 2 3 4 5

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. Savic´ & 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 Bezirks­ hauptmannschaft Spittal an der Drau (Bereich 10) for making out the per- mit, Peter Kosnik for the TLC, Othmar Breuss for the confirmation of Pla- cidiopsis oreades, and Christian Scheuer as well as Herwig Teppner for crit- ically 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. Author’s personal copy

212

6. References

Bilovitz P. O., Nascimbene J., Tutzer V. , Wallner A. & Mayrhofer H. 2014a. Terrico­ lous lichens in the glacier forefield of the Rötkees (Eastern Alps, South Tyrol, Italy). – Phyton (Horn, Austria) 54(2): 245–250. Bilovitz P. O., Tutzer V. , Wallner A., Nascimbene J. & Mayrhofer H. 2014b. Terrico­ lous lichens in the glacier forefield of the Matscherferner (Eastern Alps, South Tyrol, Italy). – Acta ZooBot Austria 150–151: 197–202. Bilovitz P. O., Wallner A., Tutzer V. , Nascimbene J. & Mayrhofer H. 2014c. Terrico­ lous lichens in the glacier forefield of the Gaisbergferner (Eastern Alps, Tyrol, Austria). – Phyton (Horn, Austria) 54(2): 235–243. Bilovitz P. O., Nascimbene J. & Mayrhofer H. 2015. Terricolous lichens in the glacier forefield of the Morteratsch glacier (Eastern Alps, Graubünden, -Switzer land). – Phyton (Horn, Austria) 55(2): 193–199. Böhm H. 1969. Die Waldgrenze der Glocknergruppe. – In: Büdel J. & Glaser U. (eds.). Neue Forschungen im Umkreis der Glocknergruppe. – Wissenschaftliche Al- penvereinshefte 21: 143–167. Breuss O. 1996. Revision der Flechtengattung Placidiopsis (Verrucariaceae). – Ös- terreichische Zeitschrift für Pilzkunde 5: 65–94. Burger R. & Franz H. 1969. Die Bodenbildung in der Pasterzenlandschaft. – In: Büdel J. & Glaser U. (eds.). Neue Forschungen im Umkreis der Glockner- gruppe. – Wissenschaftliche Alpenvereinshefte 21: 253–264 + 1 Karte. Drescher-Schneider R. & Kellerer-Pirklbauer A. 2008. Gletscherschwund einst und heute – Neue Ergebnisse zur holozänen Vegetations- und Gletscherge- schichte der Pasterze (Hohe Tauern, Österreich). – Abhandlungen der geolo- gischen Bundesanstalt in Wien 62: 45–51. Egger G. 1997. Standortsdynamik und Sukzession der Flechtenbesiedlung im Win- kelkees-Vorfeld des Seebachtales (Nationalpark Hohe Tauern, Österreich). – Wissenschaftliche Mitteilungen aus dem Nationalpark Hohe Tauern 3: 37–65. Fischer A. 2014. Gletscherbericht 2012/2013. Sammelbericht über die Gletscher­ messungen des Oesterreichischen Alpenvereins im Jahre 2013. – Bergauf 02- 2014: 34–40. Fischer M. A., Oswald K. & Adler W. 2008. Exkursionsflora für Österreich, Liech- tenstein und Südtirol, 3. Auflage – Biologiezentrum der Oberösterreichischen Landesmuseen, Linz. Frank W. 1969. Geologie der Glocknergruppe. – In: Büdel J. & Glaser U. (eds.). Neue Forschungen im Umkreis der Glocknergruppe. – Wissenschaftliche Alpen­ vereinshefte 21: 95–111. Franz H. 1969. Besiedlung der jüngst vom Eise freigegebenen Gletschervorfelder und ihrer Böden durch wirbellose Tiere. – In: Büdel J. & Glaser U. (eds.). Neue Forschungen im Umkreis der Glocknergruppe. – Wissenschaftliche ­Alpenvereinshefte 21: 291–298. Friedel H. 1951. Das Drama von Gras und Sand am Pasterzenufer. – Natur und Land 37(7–8): 124–132. Friedel H. 1956. Die alpine Vegetation des obersten Mölltales (Hohe Tauern). Erläu- terung zur Vegetationskarte der Umgebung der Pasterze (Großglockner). – Wissenschaftliche Alpenvereinshefte 16: 1–153 + 12 Tafeln + 27 Vegetation- stabellen + 2 Vegetationskarten. Author’s personal copy

213

Friedel H. 1969. Die Pflanzenwelt im Banne des Großglockners und des Pasterzen- gletschers. – In: Büdel J. & Glaser U. (eds.). Neue Forschungen im Umkreis der Glocknergruppe. – Wissenschaftliche Alpenvereinshefte 21: 233–252 + 1 Karte. Gams H. 1936. Beiträge zur pflanzengeographischen Karte Österreichs. I. Die Vege- tation des Großglocknergebietes. – Abhandlungen der zoologisch-bota­ nischen Gesellschaft in Wien 16(2): 1–79. Gams H. 1951. Die Gamsgrube an der Pasterze, das merkwürdigste „Hintergrasl“ der Alpen. – Natur und Land 37(7–8): 119–124. Hafellner J. & Türk R. 1995. Über Funde lichenicoler Pilze und Flechten im Na- tionalpark Hohe Tauern (Kärntner Anteil, Österreich). – Carinthia II 185 = 105: 599–635. Hafellner J., Muggia L. & Obermayer W. 2012. Rinodina candidogrisea, a new sore- diate species from high altitudes in the Alps. – Bibliotheca lichenologica 108: 75–102. Hartl H. 1988. Die Gamsgrube, das merkwürdigste „Hintergrasl“ der Alpen. – Carinthia II 178 = 98: 17–21. Krainer K. 1988. Ein geologischer Streifzug durch Kärnten. – Carinthia II: 178 = 98: 141–170. Krainer K. 1994. Nationalpark Hohe Tauern. Geologie – Nationalpark Hohe Tauern, Wissenschaftliche Schriften. Universitätsverlag Carinthia, Klagenfurt. Nicolussi K. & Patzelt G. 2000. Discovery of early-Holocene wood and peat on the forefield of the Pasterze Glacier, Eastern Alps, Austria. – The Holocene 10(2): 191–199. Nicolussi K. & Patzelt G. (2000) 2001. Untersuchungen zur holozänen Gletscher- entwicklung von Pasterze und Gepatschferner (Ostalpen). – Zeitschrift für Gletscherkunde und Glazialgeologie 36(1–2): 1–87. Orange A., James P. W. & White F. J. 2001. Microchemical methods for the identifica- tion of lichens. – British Lichen Society, London. Paschinger H. 1976. Kärnten. Eine geographische Landeskunde. Erster Teil: Allge- meine Darstellung. – Verlag des Landesmuseums für Kärnten, Klagenfurt. Schiechtl von H. M. & Stern R. 1985. Die aktuelle Vegetation der Hohen Tauern. Matrei in Osttirol und Großglockner. Vegetationskarten 1 : 25.000 mit Erläu- terungen. – Wissenschaftliche Schriften des Nationalparks Hohe Tauern. Universitätsverlag Wagner, Innsbruck (mit 4 Karten). Slupetzky H. 1993. Holzfunde aus dem Vorfeld der Pasterze. Erste Ergebnisse von 14C-Datierungen. – Zeitschrift für Gletscherkunde und Glazialgeologie 26(2): 179–187. Slupetzky H., Krisai R. & Lieb G. K. 1998. Hinweise auf kleinere Gletscherstände der Pasterze (Nationalpark Hohe Tauern, Kärnten) im Postglazial – Ergeb- nisse von 14C-Datierungen und Pollenanalysen. – Wissenschaftliche Mit- teilungen aus dem Nationalpark Hohe Tauern 4: 225–240. Sternberg von C. M. & Hoppe D. H. 1815. Braya, eine neue Pflanzengattung. – Denk- schriften der königlich-baierischen botanischen Gesellschaft in Regensburg: 65–75 + 1 Abbildung. Sulzer W. & Lieb G. K. 2009. Die Gletscher im Wandel der Zeit – Gletschermoni­ toring am Beispiel der Pasterze. – Vermessung und Geoinformation 3: 371– 382. Author’s personal copy

214

Tollner H. 1969. Klima, Witterung und Wetter in der Großglocknergruppe. – In: Büdel J. & Glaser U. (eds.). Neue Forschungen im Umkreis der Glockner- gruppe. – Wissenschaftliche Alpenvereinshefte 21: 83–94. Tschernutter P. 1982. Niederschläge in Kärnten 1951 bis 1980. – Schriftenreihe für Raumforschung und Raumplanung 26: 1–40 (Teil I), 1–135 (Teil II). Türk R. & Erschbamer B. 2010a. Die Flechten im Gletschervorfeld des Rotmoos­ ferners. – In: Koch E.-M. & Erschbamer B. (eds.). Glaziale und periglaziale Lebensräume im Raum Obergurgl. – Alpine Forschungsstelle Obergurgl 1: 149–157. Türk R. & Erschbamer B. 2010b. Die Flechten im Gletschervorfeld des Rotmoos­ ferners (Anhang). – In: Koch E.-M. & Erschbamer B. (eds.). Glaziale und peri­ glaziale Lebensräume im Raum Obergurgl. – Alpine Forschungsstelle Ober- gurgl 1: 278–280. Türk R. & Hafellner J. 1992. Flechten im Nationalpark Hohe Tauern – Kärntner Anteil (Österreich). – Carinthia II 182 = 102: 723–757. Türk R., Hafellner J. & Taurer-Zeiner C. 2004. Die Flechten Kärntens. Eine Be- standsaufnahme nach mehr als einem Jahrhundert lichenologischer Forschungen. – Sonderreihe Natur Kärnten 2: 1–333. – Naturwissenschaft­ licher Verein für Kärnten, Klagenfurt. White F. J. & James P. W. 1985. A new guide to microchemical techniques for the identification of lichen substances. – Bulletin of the british Lichen Society 57 (Suppl.): 1–41. Wirth V. , Hauck M. & Schultz M. 2013. Die Flechten Deutschlands. Bände 1 und 2. – Ulmer, Stuttgart. Wittmann H., Stöhr O., Pilsl P. & Gewolf S. 2009. Das Gletschervorfeld der Pas- terze (Glockner-Gruppe, Kärnten, Österreich) mit besonderer Berücksichti- gung von Ökologie, Pflanzensoziologie und Naturschutzaspekten der „alpi­ nen Schwemmlandbiotope“. – Tuexenia, Beiheft 2: 147–193. Zollitsch B. 1969. Die Vegetationsentwicklung im Pasterzenvorfeld. – In: Büdel J. & Glaser U. (eds.). Neue Forschungen im Umkreis der Glocknergruppe. – Wis- senschaftliche Alpenvereinshefte 21: 267–290 + 1 Karte.