Lichens growing on calcareous rocks in the Polish part of the Sudety
Transkrypt
Lichens growing on calcareous rocks in the Polish part of the Sudety
3 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 3 Lichens growing on calcareous rocks in the Polish part of the Sudety Mountains* MARIA KOSSOWSKA Department of Biodiversity and Plant Cover Protection, Institute of Plant Biology, University of Wroc³aw, Kanonia 6/8, 50328 Wroc³aw, Poland, e-mail: [email protected] Abstract Paper presents the results of the investigations of the calcicolous lichen flora occuring in the Sudety Mts. The study was carried out in the two most extensive areas with calcareous substrates in the Sudety Mts: the Góry Kaczawskie Mts in the western part of the Sudetes and the nie¿nik Metamorphic Region in their eastern part, and the two types of localities: natural rock outcrops and quarries. On each locality the entire lichen flora that was directly (saxicolous species) or indirectly (terricolous and muscicolous species) associated with calcareous substrates was analyzed. Of the 129 species currently found in the study area, 84 species were exclusively epilithic, 17 were epilithic, 21 species grew over mosses and 22 taxa were not connected with only one type of substrate. The calcicolous lichen flora of the Western Sudety Mts was significantly richer and more diverse than that of the Eastern Sudetes. In the Góry Kaczawskie Mts a total of 111 calcicolous lichen species were found, whereas in the nie¿nik Metamorphic Region only 72 species were identified during the study. Only 54 taxa occured in both calcareous areas in the Sudety Mts. Key words: saxicolous lichens, calciphilous lichens, Sudety Mts, Lower Silesia * The present study was partially supported by the Polish Committee for Scientific Research, grant no. 301/P04/97/13. 4 Maria-Kossowska 4 Contents Introduction . . . . . . . . . . . . . . . . . . 5 1. Study area . . . . . 1.1. Preliminary remarks . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6 1.2. Location and physiography 1.3. Geological structure . . 1.4. Climate . . . . . . 2. Materials and metods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 8 9 10 3. Historical and present-day lichen flora . . . . . 4. Ecological analysis of the present-day lichen flora . 4.1. Preliminary remarks . . . . . . . . . . . . . . . . . . . . . 14 21 21 4.2. Epilithic (saxicolous) lichens . 4.3. Epigeic (terricolous) lichens . . . . . . . . . . . . . . . . . . . . . 23 27 4.4. Epibryophytic (muscicolous) lichens . 4.5. Nitrophilous lichens. . . . . . . . . . . . . . . . . . . . . 28 29 4.6. Classification of calcareous microhabitats . . . . 5. Comparison of the calcicolous lichen flora in the Western and Eastern Sudety Mountains . . . . . . . . . 6. Threat to calcicolous lichen flora in the Sudety Mountains . . 31 . . . . 34 36 6.1. Factors of threat . . . . . . . . . . 6.2. Participation of rare and endangered species . 7. Summary and conclusions . 8. References . . . . . . 9. Streszczenie. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 37 . . . . . . . . . . . . 39 41 45 . . . . 50 104 Appendix 1. List of calcicolous lichen species found in the Polish part of the Sudety Mts . Appendix 2. Index of synonyms. . . . . . . . . . . . . . . . . . . . 5 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 5 Introduction Calcareous rocks and the soils derived from them have long been of interest to naturalists. They represent a rich and diverse habitat for calciphilous and basophilous vascular and non-vascular plants. Areas in Poland that are characterized by calcareous substrates have been the subject of a series of botanical and phytosociological monographs that deal with various systematic groups of plants and fungi. Polish lichenologists have also been studying the lichen vegetation that inhabits calcareous habitats. The present study is a continuation of efforts to fully describe the lichens of Poland and the habitats in which they are found from a geographical and ecological point of view. The geological landscape of Poland has been shaped by orogenic and glacial events. Exposed calcareous rock is found chiefly in highland and mountainous regions in the southern part of the country, including the Kraków-Wieluñ Highlands, the Pieniny Mts, the Góry wiêtokrzyskie Mts, the Western Tatra Mts, the Sudety Mts and the Przedgórze Sudeckie Foreland. Some exposed calcareous rock is found in the Wy¿yna Lubelska, Wy¿yna Ma³opolska and Wy¿yna l¹ska provinces. Furthermore, sandstones in the Beskidy Mountains usually contain varying amounts of calcium carbonate. Some of the areas mentioned have been the subject of detailed lichenological studies (Tobolewski 1958; Nowak 1961, 1967; Toborowicz 1983; Czy¿ewska 1986). In other areas, limited studies on the calcareous lichen flora have been carried out (Wójciak 1987; Kiszka & Szel¹g 1992; Kiszka 1997a, b, 2000; Donica 2001). One area that has not been the subject of study since the late 1900s and early 2000s is the Sudety Mts and their foothills in Lower Silesia. Because the natural environment has changed since then, these data are mostly of historical interest. Human activity has greatly reduced the abundance of lichens, including calcicolous taxa. There is therefore an urgent need for a detailed analysis of their distribution in the Sudety Mts and of the factors that limit their occurrence. The lichen flora that grows on calcareous substrates in the Sudetes deserve to be studied in detail also because of the specific habitat conditions found in the area. These are determined by the following factors: - Climate. The climate in the region is affected by the Atlantic oceanic airmass. This is not the case elsewhere in Poland, except in Western Pomerania in the northwestern corner of the country (Paw³owska 1972). The climate in the other calcareous areas in Poland is dominated by the continental air masses. - Substrate type. Calcareous rocks in the region are predominantly metamorphic marbles. Calcium carbonate is therefore less accessible to plants 6 Maria-Kossowska 6 (Fabiszewski 1989). This is presumably true for lichens, as well. There is also little exposed calcareous rock in the region, except in quarries. The rock surface on which lichens may grow are therefore minimally weathered. The author hopes that present study will fill in the gaps in our knowledge of the distribution of calcicolous lichens in the Sudety Mts, and, in some cases, inspire re-evaluation of their geographical and habitat preferences. Acknowledgments. The author would like to deeply thank the many people who generously provided valuable assistance during this study, including Dr. hab. Lucyna liwa, for verifying the identification of several species of the genus Lecanora, and Dr. Martin Kukwa, for identifying species of the genus Lepraria. Most of all, the author is especially grateful for the tireless attention of her advisors, Prof. Wies³aw Fa³tynowicz and Prof. Jadwiga Anio³Kwiatkowska, without whom the present work would never have been udertaken. 1. Study area 1.1. Preliminary remarks The study was carried out in the two most extensive areas with calcareous substrates in Lower Silesia: the Góry Kaczawskie Mountains in the Western Sudetes and the nie¿nik Metamorphic Region in the Eastern Sudetes. In both areas, calcareous rocks are generally crystalline limestones, and are generally exposed only in quarries. Both areas also have similar habitat conditions and a similar calcicolous lichen flora. However, the areas differ in terms of the age of the calcareous rocks, weather conditions, and surface topography. Therefore, both regions were treated separately in this chapter. The description of habitat conditions was limited to only those factors that have an immediate effect on saxicolous lichens, including geological and climatological factors. 1.2. Location and physiography nie¿nik Metamorphic Region. The name of this area comes from the geological literature. It pertains to the complex of metamorphic rocks that are exposed on the tops and slopes of the mountains in entire K³odzko region (Don & Opletal 1996). The region comprises the nie¿nik Massif including the Krowiarki Range, the Góry Bialskie Mts, and the Góry Z³ote Mts. The central part of the area represents the nie¿nik Massif proper. nie¿nik K³odzki Mt. is the highest peak in eastern part of the Polish Sudety Mountains, with an elevation of 1426 m. The Krowiarki Range is a northward ex- 7 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 7 tension of the massif, which is delineated by Puchaczówka Pass. The Krowiarki Range consists of low ridges, the highest of which is Suchoñ Mt., with an elevation of 964 m. The end of the range is marked by Wapniarka Mt. (518 m) and Dêbowa Góra Mt. (505 m) near the village of ¯elazno. The Góry Bialskie Mts are an eastward extension of the nie¿nik Massif, and are delineated by the P³oszczyna Pass. The highest peak in the range is Postawna Mt., with an elevation of 1124 m. They have been included in the Góry Z³ote Mts mesoregion (Kondracki 1994). The Góry Bialskie Mts consist of a low, continuous massif divided by deep gorges into several individual ridges. They are separated from the Góry Z³ote Mts proper by the U Trzech Granic Pass and the valley of Górna Bia³a L¹decka river. The Góry Z³ote Mts are one of the longest ranges in the Sudetes. They begin in Poland at the Prze³êcz K³odzka Pass, which separates them from the Góry Bardzkie Mts. They then continue southeastward into the Czech Republic to the Ramzowska Pass, which divides them from the Hrubý Jeseník Massif. To the north, the Góry Z³ote Mts are delineated by the edge of the Sudetic Marginal Fault. The landscape of the entire nie¿nik Metamorphic Region is characterized by rounded mountain ridges divided by deep gorges. There are exposed few rock formations that provide a natural habitat for epilithic lichens. There are, however, many abandoned quarries in the region in which rocks of various types are exposed and colonized by lichens. Góry Kaczawskie Mountains. This range is part of the Western Sudetes, together with the nearby Karkonosze Mts, the Góry Izerskie Mts and the Rudawy Janowickie Mts. The Góry Kaczawskie Mts are relatively low. The highest peak in the range is Skopiec Mt., with an elevation of 728 m. The Góry Kaczawskie Mts are extensive. On the south, they border with the Kotlina Jeleniogórska Basin. On the west, they are delineated by the valley of Bóbr river, which separates them from the Góry Izerskie Mts. On the east, they border with the Pogórze Bolkowskie Foothills in the Central Sudetes. On the north, they blend into the Pogórze Kaczawskie Foothills, which lie on a separate tectonic fragment (Kondracki 1994). The Góry Kaczawskie Mountains proper consist of several more or less parallel sub-ranges that run from northeast to southwest. They are divided by the valley of Kaczawa river into two unequal parts. The larger western part (called also the Western Ridge) consist of the Southern Ridge, the Northern Ridge, and the Small Ridge. The Southern Ridge is the highest in the entire Góry Kaczawskie Mts. It is separated from the Northern Ridge by the valleys of Lipka and wierzawa. The smaller eastern part (the Eastern Ridge) is delineated by the valleys of Kaczawa and Nysa Szalona rivers. 8 Maria-Kossowska 8 Although the Góry Kaczawskie Mts are relatively low, they are very diverse in terms of landscape. This reflects the complex geological understructure on which they lie, which is composed of many different types of rock. The area is characterized by deep, twisted valleys and steep hill slopes. There are numerous exposed rock formations in the area, as well as many quarries, including limestone quarries, some of which are extensive. 1.3. Geological structure nie¿nik Metamorphic Region. In the Eastern Sudety Mountains, calcareous rocks are usually associated with the nie¿nik Metamorphic Region, the complex of ancient metamorphic rocks that includes the nie¿nik Massif proper, the Góry Bialskie Mts and the Góry Z³ote Mts. The rock formations in the area date back as far as the Precambrian, when the seabed that existed here at the time was covered by deposits of sands, limestones, muds and slates. Local deposits of basaltic lavas and tuffs were results of underwater volcanic activity. This diverse collection of sedimentary and igneous rocks is called the Stronie Series (Don & Opletal 1996). During later metamorphosis, the crystalline structure of the Stronie Series changed, producing paragneisses, micaschists, quartzites, marbles and amphibolites. Together with orthogneisses, these rocks are found exposed on tops and slopes of the the Eastern Sudetes. The rock formations of the Stronie Series reach the surface on about half of the area of the nie¿nik Metamorphic Region. They are common in the Krowiarki Range, in the Góry Z³ote Mts and near Stronie l¹skie. The fundamental components of the series are various mica-schists interspersed with numerous lenticular formations of crystalline limestone and dolomite marbles that become erlans where they come in contact with nie¿nik gneisses. The marbles sometimes form layers up to several hundred meters in thickness. The largest lenticular formations of marbles are found in the Krowiarki Range and near Stronie l¹skie, Kletno and Z³oty Stok. These rocks are very rarely exposed as natural rock outcrops. One exception is the rock Pulinka on the northeast slope of ¯mijowiec Ridge, above the quarry designated Kletno II. Another is the rock Kapelusz on the northwest slope of Dêbowa Góra Mt. in the Krowiarki Range. Góry Kaczawskie Mountains. Limestones and dolomites in the Góry Kaczawskie Mts are associated with an early Paleozoic complex of metamorphic rocks called the Góry Kaczawskie Metamorphic Region. This region extends throughout the entire range of the Góry Kaczawskie Mts into the surrounding areas, including the southern part of the Pogórze Kaczawskie Foothills and the eastern part of the Pogórze Izerskie Foothills. This complex consists not only of crystalline limestones and dolomites, but also of a series of metamor- 9 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 9 phic rocks of different origin and mineral content, including phyllites, greenstones, greenstone schists and sericite schists. Many of the calcareous rocks in the Góry Kaczawskie Mts are from the lower and middle Cambrian, when the region was a sea basin in which calcareous sediments were deposited. Previously, silt and mud had been deposited, together with intermittent layers of acidic and basic lava. The series of calcareous rock was also later covered by formations of igneous origin (Kozdrój 1995). The entire complex of sedimentary and igneous rocks subsequently underwent metamorphosis during the Hercynian Orogeny, when the folding that formed the Góry Kaczawskie Mts took place. In the process, the calcareous deposits became crystalline limestones of the type called Wojcieszowskie limestones. In the Western Sudetes, the area in which crystalline limestones occur can be divided into two zones (Sawicki 1979). The western one extends from Jêdrzychowice near Zgorzelec to Siedlêcin near Jelenia Góra. It covers the entire area of the Pogórze Izerskie Foothills and the western part of the Góry Kaczawskie Mts. The area is characterized by calcareous rock that occurs in small, isolated lenticular formations. The eastern zone extends from Dziwiszów eastward to Lipa and Pogwizdów. Crystalline limestones and dolomites form close, sometimes bipartite layers of considerable thickness that stretch uninterrupted over 18 km. In comparison to the Precambrian marbles of the Stronie Series found in the Eastern Sudetes, the limestones of the Góry Kaczawskie Mts are less metamorphosized and can be described as epi-metamorphic. They are mainly varieties composed of small or very small crystals. Varieties composed of large crystals called sugar marbles occur only locally. In terms of mineralogical composition, the limestones of the Góry Kaczawskie Mountains represent a complex of interlacing calcareous and dolomitic rocks with varying amounts of calcite and dolomite. In the Góry Kaczawskie Mts, exposed calcareous rocks are found in natural formations on hilltops and hillsides, and also in the many quarries in the area. 1.4. Climate The climate of the Sudety Mountains is shaped by many factors, the most important of which are: latitude, longitude, altitude and surface topography. These factors determine basic meteorological parameters such as sunlight, temperature, relative humidity, precipitation and wind direction. In accordance with the classification of climatic types in Poland proposed by Romer (1949), the Sudety Mts belong to the submontane/montane region, which is characterized by low temperatures at higher altitudes, cool summers, 10 Maria-Kossowska 10 abundant precipitation, frequent strong winds, and a small amplitude in annual temperature (Medwecka-Korna 1972). The climate in the Sudety Mts is markedly milder than that of the Carpathians to the east because of the influence of the Atlantic oceanic airmass, which is not impeded by any north-south running mountain ranges to the west (Wo 1995). These effects manifest themselves as warmer thermal isoanomalies, gentler oscillations in temperature, more abundant precipitation and more frequent fog cover than in the Karpaty Mountains (Medwecka-Korna 1972). Their effect is strongest in the Góry Kaczawskie Mts at the western end of the Sudetes, and weakest in the nie¿nik Metamorphic Region at the eastern end. In accordance with the map of climatic regions in the Sudety Mts prepared by Schmuck (1960), the areas studied belong to two different pluvio-thermal regions characterized by different patterns in annual temperature and precipitation. The mean temperatures and total annual precipitation for the areas is compared in Table 1. Table 1. Selected climatic features of the Western and Eastern Sudety Mts Mean temperature [oC] Precipitation [mm] Western Sudety Mts 5–7 650 – 850 Eastern Sudety Mts 3–8 600 – 1000 Region Based on data from 1950 to 1981; source: B¹c-Bronowicz 1997, Piasecki 1997. In the entire Sudety Mts, prevailing winds are westerly. During winter, they are mostly southwesterly, and during summer, northwesterly (Schmuck 1969). Local wind patterns may depend on the surface topography and the presence of nearby mountain massifs. From 1966 to 1985, winds were generally westerly in the Góry Kaczawskie Mts, and generally southerly in the nie¿nik Metamorphic Region (Lorenc 1997). 2. Materials and methods The present analysis of the calcicolous lichen flora of the Sudety Mountains was based chiefly on materials collected from the field from 1996 to 1998. Historical data came from earlier studies carried out by Flotow (1850, 1851), Körber (1855) Stein (1879, 1889), Eitner (1896, 1901, 1911) and Fabiszewski (1968). The material included also specimens housed at the herbaria of the University of Wroc³aw (WRSL) and the University of Poznañ (POZ). 11 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 11 During field studies, only material growing on exposed natural calcareous rock was collected. No specimens were included that were found growing on concrete, mortar and other anthropogenic substrates. Material was collected from a total of 45 objects (localities): rocks, outcrops and quarries. They are listed and mapped below. Within each locality, lichens were collected at several points, each representing a different habitat type. They are called collection sites. Because the boundaries between the habitat types were not clearly delimited, analysis was carried out on the total lichen flora growing on calcareous substrates at each locality. The lichens collected included speciemens connected directly with rock substrate (epilithic) and indirectly (epigeic and epibryophytic). The collected material was submitted to the herbarium of the University of Wroc³aw (WRSL). The material collected was determinated using keys for identifying lichens of Nowak & Tobolewski (1975) Wirth (1980, 1995) Purvis et al. (1992) and Ozenda & Clauzade (1970), as well as monographs on individual taxonomic groups, e.g. Degelius (1954), Servít (1954) and Jorgensen (1994). Lichens of / the genus Lepraria were identified using thin layer chromatography (TLC). The nomenclature used follows Hafellner and Türk (2001), Fa³tynowicz (2003) and Santesson et al. (2004), with only exceptions for the genus Bilimbia (Veldkamp 2004) and Xanthoria s.l. (Sochting et al. 2002). / In the analysis of present-day lichen flora a five-point scale of frequency was used. The scale is based on proporcional share of species in a total number of localities (Tab. 2). Table 2. The species frequency scale Share [%] Number of localities Common Category 75 – 100 34 – 45 Very frequent 50 – 75 23 – 33 Frequent 33 – 50 15 – 22 Dispersed and rare 5 – 33 3 – 14 <5 1–2 Very rare The distribution of each taxon was mapped in accordance with a modificated ATPOL grid square system (Zaj¹c 1978, Cieliñski & Fa³tynowicz 1993). The basic unit was a square with sides 10 km long. Mapping was carried out using the Gnomon 3.2 software (Desmodus, Poland). 12 Maria-Kossowska 12 List of localities Te localities are arranged in accordance with ATPOL system. Numbering of localities is consistent with the Figure 1. Fig. 1. Distributin of the localities in the ATPOL grid squares Ea59 Pogórze Izerskie Foothills, Wzgórza Radomickie Hills: (1) Wapienica rise about 0.5 km east of Radomice, abandoned quarry (elev. about 460 m); (2) eastern slope-foot of the Wapniak hill (391 m) near Pilchowice, small break with a southern exposure (elev. about 310 m); (3) northeast slope of Wapniak hill near Pilchowice, little rock in the forest (elev. about 330 m). Ea69 Góry Kaczawskie Mts, Western Ridge: (4) Wapienna Mt. (508 m) north of Siedlêcin, abandoned quarry at the top of the mountain (elev. about 480 m). 13 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 13 Eb60 Góry Kaczawskie Mts, Western Ridge: (5) Kapela rise between Dziwiszów and Podgórki, old quarry on the eastern slope (elev. about 590 m); (6) Kapela rise, active quarry with unexploited, north-facing wall (elev. about 585 m); (7) Kapela rise, low rocky crest in xerothermic grassland above an active quarry (elev. about 600 m); (8) Kapela rise, small break in xerothermic grassland (elev. about 600 m); (9) Bia³e Ska³y rocks on the northern slope of Malak Mt. (720 m) above Podgórki (elev. about 560 m); (10) Piwniczna rock on the northern slope of Malak Mt. (elev. about 620 m). Eb61 Góry Kaczawskie Mts, Western Ridge: (11) Podgórki, extensive quarry 1 km north of the village (elev. about 445 m). Góry Kaczawskie Mts, Eastern Ridge: (12) Wojcieszów, Gruszka quarry at the western footslope of Chmielarz Mt. (581 m), in the centre of the town (elev. about 450 m); (13) Mi³ek Mt. (594 m) near Wojcieszów, rock ledge on the top of M³yniec peak (the northern summit of Mi³ek Mt.; elev. about 565 m); (14) Mi³ek Mt., west-facing vertical outcrop below M³yniec peak (elev. about 550 m); (15) Mi³ek Mt., outcrop on the ski slope with north-eastern exposure (elev. about 480 m); (16) Mi³ek Mt., abandoned quarries at the south-western footslope (elev. about 475 m); (17) Mi³ek Mt., rocks on the top of Cisowa peak (the southern summit of Mi³ek Mt.; elev. about 595 m); (18) Mi³ek Mt., steep rock on the south-eastern slope of Cisowa peak (elev. about 570 m); (19) Ose³ka Mt. (591 m) about 0,5 km north-east of Mys³ów, abandoned quarry on the eastern slope (elev. about 510 m); (20) Stary Wapiennik rise east of Mys³ów, active quarry with unexploited walls facing south and east (elev. about 485 m). Pogórze Kaczawskie Foothills, Pogórze Wojcieszowskie Foothills: (21) Lipa, small quarry among cultivated fields, about 1 km west of the village (elev. about 435 m); (22) Lipa, rock outcrops on a low hill west of the village (elev. about 450 m). Eb62 Pogórze Kaczawskie Foothills, Pogórze Wojcieszowskie Foothills: (23) Bukowa Hill (431 m) between Jastrowiec and Grudno, abandoned quarry at the northern footslope (elev. about 380 m); (24) Wapniki Hill (478 m), extensive quarry on the southern slope (elev. about 480 m); (25) Wapniki Hill, small break at the the eastern footslope (elv. about 425 m); (26) Nowe Rochowice, high quarry north-east of the village, ca. 400 m from the Bolków Jelenia Góra road (elev. about 470 m); (27) Rochowicka Ska³a rock on the eastern edge of Nowe Rochowice (elev. about 425 m). Fb26 nie¿nik Massif, Krowiarki Range: (28) Wapniarka Mt. (518 m) about 1.5 km south of ¯elazno, abandoned, deep quarry at the north-east- 14 Maria-Kossowska 14 ern footslope (elev. about 430 m); (29) Wapniarka Mt., abandoned quarry on the south-eastern slope above the previous locality (elev. about 475 m); (30) Dêbowa Góra Mt. (506 m) about 1.5 km south of ¯elazno, overgrown quarry near the pass between Wapniarka Mt. and Dêbowa Góra Mt. (elev. about 425 m); (31) Dêbowa Góra Mt., abandoned quarry on the north-eastern slope (elev. about 400 m). Fb27 Góry Z³ote Mts: (32) Z³oty Stok, abandoned quarry in the northern part of the town (elev. about 340 m). . Fb36 nie¿nik Massif, Krowiarki Range: (33) Wapniarka Mt. (518 m), south of ¯elazno, rock outcrops on escarp at the south-western footslope (elev. about 460 m); (34) Wapniarka Mt., small break close by previous locality; (35) Wapniarka Mt., rocks at the top of the mountain (elev. about 515 m); (36) S³upiec Mt. (531 m) south of O³drzychowice K³odzkie, quarry on the western edge of the ridge, by K³odzko Bystrzyca K³odzka road (elev. about 385 m); (37) S³upiec Mt., heavily overgrown quarry north of the previous locality (elev. about 390 m); (38) Sêdzisz Mt. (489 m) in the S³upiec ridge south of O³drzychowice K³odzkie, active quarry (elev. about 395 m); (39) Krzy¿owa Mt. (508 m) about 2 km north-east of Stary Waliszów, remains of a quarry in a top part of the mountain (elev. about 500 m). Fb37 Góry Z³ote Mts: (40) Jaskinia Radochowska cave at the southern footslope of Bzowiec Mt. (697 m) about 2 km north-west of Radochów, rock face near the entrance to the cave (elev. about 365 m). nie¿nik Massif, Krowiarki Range: (41) Rogó¿ka, quarry at the southwestern footslope of Wapnisko Mt. (799 m; elev. about 730 m). Fb47 nie¿nik Massif, Dolina Klenicy valley: (42) Kletno, abandoned quarry Kletno II at the eastern footslope of ¯mijowiec ridge (1153 m), ca. 1 km south of the last buildings in the village (elev. about 800 m); (43) Kletno, Pulinka rock on the eastern slope of ¯mijowiec ridge, above the quarry Kletno II (elev. about 880 m); (44) Kletno, rock outcrop at the entrance to Jaskinia Niedwiedzia Nature Reserve (elev. about 750 m); (45) Kletno, rock face behind the entrance pavillion to Jaskinia Niedwiedzia cave at the western footslope of Stroma Mt. (1167 m; elev. about 810 m). 3. Historical and present-day lichen flora The calcicolous lichen flora of the investigated area (174 taxa; Tab. 3) consists of two fundamental categories: 15 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 15 - present-day lichen flora lichens found during the field studies of 1996 1998. - historical lichen flora lichens recorded before 1970, mentioned in publications or kept in lichen herbaria. Data on historical lichen flora were gathered primarily from the reports of surveys carried out from the 1800s and early 1900s. The most valuable source was the monograph published by Stein (1879), which summarized the current state of knowledge on the lichen flora of Silesia, on the basis of both the authors personal observations as well as on data from previous studies such as those of Flotow (1850, 1851) and Körber (1855, 1865). This monograph was later supplemented by the author himself in1889, and then by Eitner in 1896, 1901 and 1911. After the Second World War, lichenological investigations of calcareous substrates in the Sudety Mts were carried out by Fabiszewski. In his monograph on epiphytic lichen communities of the nie¿nik Massif (1968) he also noted the presence of several calcicolous lichens at five of the study localities. Since then, a survey was carried out by Tobolewski on Mi³ek Mt. in the Góry Kaczawskie Mts. He included some of the data in his Atlas of the Geographical Distribution of Lichens in Poland (Tobolewski 1979, 1980). The historical lichen biota of investigated area contains 108 certain species (localities of which were precisely described). Another 13 species are probable, because they were reported by Stein (op. cit.) as either abundant or common in Silesia as a whole, without any localities. However, only those species from the latter group that are still found in the study area were included in the present study. Of the 108 species mentioned in the historical literature, 45 are currently not found in the study area. They may have disappeared because of habitat degradation caused by quarrying, or because of changes in habitat conditions such as the shading by trees of previously exposed rock formations. Human activity has played a key role in the disappearance of some species. Air pollution in particular has had a negative impact on calcicolous lichens in the area, though not as great an effect as it has had on epiphytic lichens. Some species may have been simply overlooked in recent surveys because of their small size. This may be true for example for species of the genera Thelidium and Verrucaria. It is not unlikely that some of these species will be found during more thorough surveys in the future. The occurence of other 63 species mentioned in the historical literature was confirmed during the study. 21 of them have been found at at least one site mentioned in the literature: Aspicilia calcarea, A. contorta, A. coronata, Caloplaca chalybaea, C. cirrochroa, C. variabilis, Candelariella aurella, Catillaria lenticularis, Clauzadea monticola, Collema fuscovirens, 16 Maria-Kossowska 16 Dermatocarpon miniatum, Lecidella stigmatea, Leptogium gelatinosum, Protoparmeliopsis muralis, Rhizocarpon petraeum, Sarcogyne regularis, Thelidium gisleri, Verrucaria hochstetteri, V. muralis and V. nigrescens. Most of them were found also at other, new localities. Only two species, Aspicilia coronata and Rhizocarpon petraeum, were found at single localities in the past and are still found only on these sites. The present-day lichen flora of the study areas is represented by 129 species. This includes certain historical species confirmed during the study (see above), species probable which might be collected here in the past, as well as 53 new species which had not been previously reported as growing in the study area. Most of them had never been previously found in the Sudety Mts, and were also uncommon or rare in the rest of Poland. A few had been found previously in the Sudetes, but only on completely different types of substrate (see chapter 4.2). These species can therefore be considered to be new for limestones of the Sudety Mts. The proportion of particular categories in total calcicolous lichen flora of the investigated area is shown on diagram (Fig. 2). 78% of the 174 species included in this study are currently found in the study area, whereas 22% are no longer found (Fig. 2). These proportions are typical for areas strongly affected by human activity (Cieliñski & Czy¿ewska 1992). In addition, most of the species currently found in the area were found at only one locality. This indicates that the number of species growing in the study areas will probably decrease in the future. 0% 20% 40% 1 60% 2 80% 3 100% 4 Fig. 2. Participation of historical and presently occuring species in the investigated lichen flora 1 exclusively historical species; 2 confirmed historical species; 3 'probable' historical species; 4 new species 17 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains Table 3. Historical and present-day lichen flora of the investigated area. Species 1 Acarospora cervina Acarospora glaucocarpa Acarospora heppii Acarospora macrospora Acrocordia conoidea Agonimia gelatinosa Arthonia lapidicola Arthopyrenia inconspicua Aspicilia calcarea Aspicilia contorta subsp. contorta Aspicilia contorta subsp. hoffmanniana Aspicilia coronata Aspicilia farinosa Aspicilia moenium Bacidia bagliettoana Bacidia trachona Bacidina egenula Bilimbia lobulata Bilimbia sabuletorum Caloplaca alociza Caloplaca cerina var. cerina Caloplaca cerina var. muscorum Caloplaca chalybaea Caloplaca cirrochroa Caloplaca citrina Caloplaca coronata Caloplaca crenulatella Caloplaca decipiens Caloplaca erythorcarpa Caloplaca holocarpa Caloplaca lactea Caloplaca neglecta Caloplaca saxicola Caloplaca teicholyta Caloplaca variabilis Caloplaca velana Candelariella aurella Candelariella vitellina Catapyrenium cinereum Report till 1945 1945 – 1970 2 3 + + + + + + + + + + + + + + + + ? + + + + + + + + + + + + + + + + + + + + + + + + + + + + + ? + ? 4 + + + + + + + + + present + + + + + + + 17 18 Maria-Kossowska 1 Catapyrenium daedaleum Catillaria chalybeia Catillaria lenticularis Cladonia glauca Cladonia pocillum Clauzadea monticola Collema auriforme Collema callopismum Collema crispum Collema cristatum Collema fuscovirens Collema limosum Collema polycarpon Collema tenax Dermatocarpon miniatum Diploschistes gypsaceus Diploschistes muscorum Diplotomma alboatrum Eiglera flavida Endocarpon adscendens Endocarpon pusillum Farnoldia hypocrita Farnoldia jurana Fulgensia fulgens Gyalecta jenensis Heppia lutosa Hymenelia epulotica Hymenelia prevostii Lecania erysibe Lecanora albescens Lecanora campestris Lecanora crenulata Lecanora dispersa Lecanora flotowiana Lecidella carpathica Lecidella stigmatea Lemmopsis arnoldiana Lempholemma chalazanum Lempholemma polyanthes Lepraria crassissima Lepraria eburnea Lepraria lobificans Lepraria vouauxii 2 18 3 4 + + + + + + + + + + ? + + + + + + + + + + ? ? ? + + + + + + + + + + + + ? ? + + + + ? + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + 19 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 1 Leptogium gelatinosum Leptogium intermedium Leptogium lichenoides Leptogium plicatile Leptogium schraderi Leptogium subtile Leptogium tenuissimum Lobothallia radiosa Megaspora verrucosa Mycobilimbia lurida Mycobilimbia pilularis Mycobilimbia tetramera Opegrapha calcarea Opegrapha dolomitica Opegrapha rupestris Peccania coralloides Peltigera didactyla Peltigera praetextata Peltigera rufescens Phaeophyscia nigricans Phaeophyscia orbicularis Phaeophyscia sciastra Physcia adscendens Physcia caesia Physconia grisea Placidium rufescens Placidium squamulosum Placynthium nigrum Placynthium tremniacum Polyblastia albida Polyblastia dominans Polyblastia sepulta Polyblastia theleodes *Polycoccum marmoratum Porocyphus rehmicus Protoblastenia incrustans Protoblastenia rupestris Protoparmeliopsis muralis Pseudosagedia byssophila Psorotichia schaereri Rhizocarpon petraeum Rhizocarpon umbilicatum Rinodina bischoffii 2 3 + ? + + + + + + + 4 + + + + + + + + + + + + + + ? ? ? + + + + + + + + + + + + + + + + + + + + + + ? ? + + + + + + + + + + + + + + 19 20 Maria-Kossowska 1 Rinodina teichophila Sarcogyne clavus Sarcogyne regularis Sarcopyrenia gibba Scoliciosporum umbrinum Solorina saccata Squamarina cartilaginea Squamarina lentigera Staurothele guestphalica Staurothele hymenogonia Staurothele succedens Synalissa symphorea Thelidium aphanes Thelidium cataractarum Thelidium decipiens Thelidium gisleri Thelidium incavatum Thelidium papulare Thelidium zwackhii Thelocarpon impressellum Thermutis velutina Toninia aromatica Toninia candida Toninia sedifolia Toninia subnitida Verrucaria apomelaena Verrucaria caerulea Verrucaria cincta Verrucaria deminuta Verrucaria dolosa Verrucaria elaeomeleana Verucaria fuscella Verrucaria hochstetteri Verrucaria infumata Verrucaria longicollis Verrucaria muralis Verrucaria murina Verrucaria nigrescens Verrucaria nigroumbrina Verrucaria obfuscans Verrucaria procopii Verrucaria saprophila Verrucaria sylvatica 2 20 3 4 + + + + + + + + + + + + + + + + + + + + + + + + + + + + ? + + + + + + + + + + ? + + + + + ? + + + + + + + + + + + + + 21 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 1 Verrucaria viridula Xanthomendoza fallax Xanthoria calcicola Xanthoria candelaria Xanthoria elegans Xanthoria parietina 2 3 + 21 4 + + + + ? + + + Key: + certain species; ? probable species (see text) 4. Ecological analysis of the present-day lichen flora 4.1. Preliminary remarks The species composition of lichen vegetation growing in a given locality depends on several ecological factors, including the chemical nature of the substrate. For calcicolous lichens, the pH and calcium carbonate content of the substrate are of key importance. Other factors that affect the lichen flora include: - availability and intensity of sunlight and shade; - humidity and the moisture content of the substrate; - exposure and weathering of the substrate; - interactions with other organisms such as mosses and vascular plants. As it was described in detail in chapter 2, the present study took into account the entire lichen flora that was directly or indirectly associated with calcareous substrates at the localities studied. The species identified greatly differed in terms of habitat requirements. Based on substrate preference, the species identified can be classified as epilithic, epigeic or epibryophytic. The number of species found on each type of substrate are presented in Figure 3. The distinctions between these classes are not always clear. Some species are found only on one particular type of substrate (exclusive species). Some occasionally grow on another type of substrate in addition to their primary substrate (passing species). Other taxa are found in approximately equal abundance on two different types of substrate. For example, Physcia adscendens and Xanthoria parietina were found both on mosses growing on rock surfaces, as well as on the rock surfaces themselves. Three others, Aspicilia contorta subsp. contorta, A. moenium and Protoblastenia rupestris are epilithic species that were occasionally found growing on soil. Of the 129 species currently found in the study area, 97 were epilithic. Of these, 84 species were exclusively epilithic. 33 species were epigeic, of which 17 were exclusive epigeic. 21 species were epibryophytic, of which only six were exclusively epibryophytic. The rest were also found on rock surfaces or soil as well. 22 Maria-Kossowska 22 1 M 2 T S 0 10 20 30 40 50 60 70 80 90 100 Fig. 3. Number of species in particular ecological groups of the investigated lichen flora S epilithic species, T epigeic species, M epibryophytic species; 1 exclusive species, 2 'passing' species. 4.2. Epilithic (saxicolous) lichens General characteristics. Of the 97 epilithic species growing directly on rock surfaces, almost 90% was exclusively epilithic (see chapter 4.1). The other 10% made a group of passing species. Among them, three were also occasionally found on soil near rock surfaces, four species were lichens typical for other types of substrate, which were found exceptionally on rock surfaces and six species have no clear substrate preference (Tab. 4). Table 4. Epilithic passing species growing on various types of substrates, including soil (s) amd mosses (m) Epilithic species exceptionally found on other substrates Species of other substrates exceptionally found on rocks Aspicilia contorta subsp. contorta (s) Aspicilia moenium (s) Protoblastenia rupestris (s) Bilimbia sabuletorum (m) Diploschistes muscorum (m) Leptogium intermedium (m) Collema crispum (s) Habitat non-sepcific species Lepraria eburnea (s, m) Lepraria lobificans (s) Lepraria vouauxii (s, m) Physcia adscendens (m) Xanthoria candelaria (m) Xanthoria parietina (m) 23 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 23 The epilithic species in the study area are very diverse in terms of their frequency (Fig. 4). Frequent and common species, which are the bulk of each lichen flora and determine its nature, made up only slightly more than 9% of the epilithic lichen flora in the study area. They are (in brackets the numbers of localities): Aspicilia contorta subsp. contorta (19), Caloplaca citrina (15), Lecanora albescens (18), Lecanora dispersa (31), Lecidella stigmatea (23), Sarcogyne regularis (24), Protoblastenia rupestris (33) i Verrucaria nigrescens (15). These species occure throughout Poland, and are not selective in terms of habitat requirements. They are usually found not only on naturally occurring rock surfaces, but on concrete and mortar as well (see S¹gin 1998). 0% 20% 40% 1 60% 2 80% 3 100% 4 Fig. 4. Participation of saxicolous lichen species in particular frequency classes 1 very rare species, 2 rare and dispersed species, 3 frequent species, 4 very frequent species The richness of epilithic lichen flora in the study area was chiefly determined by very rare species. Each was found at only one or two sites in the study area. They include: Acarospora macrospora, Arthonia lapidicola, Arthopyrenia inconspicua, Aspicilia coronata, A. farinosa, Caloplaca teicholyta, C. cerina var. cerina, C. coronata, C. saxicola, Candelariella vitellina, Collema callopismum, C. crispum, Diploschistes gypsaceus, D. muscorum, Farnoldia hypocrita, Hymenelia epulotica, H. prevostii, Lecania erysibe, Lecanora campestris, Lemmopsis arnoldiana, Leptogium intermedium, Phaeophyscia sciastra, Polyblastia albida, P. theleodes, Protoblastenia incrustans, Psorotichia schaereri, Rhizocarpon petraeum, Rinodina teichophila, Sarcogyne clavus, Sarcopyrenia gibba, Scoliciosporum umbrinum, Staurothele hymenogonia, Thelidium cataractarum, 24 Maria-Kossowska 24 T. incavatum, T. papulare, Thermutis velutina, Toninia subnitida, Verrucaria fuscella, V. procopii, V. sylvatica, V. viridula, Xanthomendoza fallax and Xanthoria candelaria. These species made up approximately 50% of the total epilitic lichen flora in the study area. They include species that are rare in Poland as a whole, as well as species that are common elsewhere, such as Caloplaca saxicola and Lecania erysibe. In addition, a distinct group of species usually found on other types of substrate can be separated. For example, some are typical for silicaceous rocks, such as Catillaria chalybeia, Lecanora campestris, Lecidella carpathica, Sarcogyne clavus and Scoliciosporum umbrinum. The pattern of epilithic lichen flora shown here, which is predominantly made up of very rare species, and only partially made up of frequent and common species, is typical for regions with extremely diverse ecological factors and dispersed potentially suitable habitats. On the other hand, it may indicate intensive extinction process on the study area species that were once common are now restricted to scattered, isolated localities that provide exceptionally good habitat conditions. Share of morphological forms. Apart from characteristic species composition mentioned above, epilithic lichen fora connected with calcareous rocks is distinguished by characteristic combination of morfological types of thalli. Contrary to siliceous substrates, on which foliose lichens belonging to genera such as Umbilicaria i Parmelia s. l. dominate, calacreous rocks are colonized predominantly by crustose species that are often endolithic. Only a few of calcicolous lichens growing on rocks have foliose thalli (mostly species of the family Collemataceae). Fruticose species are practically never found (see Fröberg 1989; S¹gin 1998). This typical for calcareous areas pattern was also largely true for the areas included in the present study. Six morphological forms of epilithic lichens were identified in the study area (Fig. 5): crustose epilithic lichens (cep) thalli superficial, either well developed or almost inapparent, continuous to areolate. In the study area, there were 63 species in this group, and they made up the bulk of the lichen flora. crustose endolithic lichens (cen) or with endolithic hypothallus (cep/n) thalli either totally or partially immersed in upper layers of a rock Most of them form perithecia, e.g. species from genera Verrucaria, Thelidium, Polyblastia and Sarcopyrenia. In the study area, there were 12 species in this group, including three that are totally endolithic. Only one endolithic species that forms apothecia was found: Protoblastenia incrustans. 25 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 0% 20% cep 40% cen i cep/n 60% plac 80% sq fol 25 100% fil Fig. 5. Participation of particular morphological types of thalli in saxicolous lichen flora (see text) placodioid lichens (plac) intermediate form between crustose and foliose thalli. Central part of thalli is crustose, with distinctly radiating marginal lobes. In the study area, there were four species in this group: Caloplaca cirrochroa, C. decipiens, C. saxicola and Protoparmeliopsis muralis. squamulose lichens (sq) a small group of lichens sometimes treated together with crustose ones. In the study area, there were two species in this group: Aspicilia moenium and Leptogium intermedium. foliose lichens (fol) thalli relatively loosely connected to the substrate by holdfasts or rhizines. In the study area, there were 14 species in this group. About a third of them were cyanobacterial species belonging to the genera Collema and Leptogium of the family Collemataceae. Greenalgal lichens included Dermatocarpon miniatum and species belonging to the genera Physcia, Phaeophyscia and Xanthoria which were generally found in habitats rich in nitrogen compounds. filamentous lichens (fil) In the study area, there was one species included in this group: Thermutis velutina, as lichen forming pillows or tufts of filamentous photosynthetic symbiont (Scytonema) interwoven with fungal hyphae. Of these groups, the crustose endolithic and semi-endolithic (with endolithic hypothallus) species are particularly noteworthy. In spite of the rule presented above, species of this group made up only 17.7% of the total lichen flora in the areas examined. The group share is actually much smaller if these species are excluded that have a well-developed superficial thallus and only an 26 Maria-Kossowska 26 endolithic hypothallus. This feature is typical for the Sudety Mts, and sets them apart from other calcareous regions in Poland. It is connected with the specific properties of the calcareous substrate in the region. The sedimentary rocks that once underlay the region have undergone metamorphic process, and have become very dense and hard crystalline limestones. The calcium carbonate crystals of which they are formed are relatively insoluble. It is very difficult for endolithic lichens to colonize this kind of substrate. A similar phenomenon was observed on hard Ordovician limestones in southern Scandinavia (Fröberg 1989). Physiological types of calcicolous lichens. The term calciphilous lichens is often applied to all species that are found on calcareous substrates. In reality, this group is very heterogenous, and various physiological types can be distinguished: calciphilous lichens sensu stricto, basophilous lichens which prefer basic or neutral substrates and may also be found on basic siliceous rocks such as basalts, gabbros, greenstones and amphibolites, ubiquitous lichens and nitrophilous species, which prefer habitats rich in fixed nitrogen, both calcareous and non-calcareous, regardless of the pH of the substrate. The group of calciphilous lichens sensu stricto which require calcium carbonate for their physiological activites may also be heterogenous, since it is not completely known if and for which species the presence of Ca++ or CO3-- ions is significant. Many strong calcicoles (according to Gilbert 1993), such as Aspicilia contorta subsp. hoffmanniana, Caloplaca lactea, Lecanora albescens and Peccania coralloides, occure also on gypsum (hydrated calcium sulphate CaSO4×2H2O) in the Niecka Nidziañska Basin (Nowak 1974, Kiszka 1987). This seems to confirm at least partial dependence of this group on calcium ions. Further laboratory studies are needed to resolve the question. In the investigated area in the Sudety Mts, the calcicolous lichen flora is also heterogenous and is probably made up of representatives of all of the physiological types mentioned above. There is also a distinct group of species that are typically associated with siliceous substrates. This group included Catillaria chalybeia, Lecanora campestris, Lecidella carpathica, Sarcogyne clavus, and Scoliciosporum umbrinum. It is difficult to explain why they were found growing on calcareous substrates in the study area. It may be connected with local mineralizations, which sometimes occur in crystalline limestones. Weathering of such rocks may cause local acidification of the rock surface (Gilbert 1993). The occurence of silicicolous species is the characteristic trait of calcicolous lichen flora in the Sudety Mts, as they have never been reported from other calcareous regions in Poland. 27 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 27 It is also difficult to separate calciphilous lichens sensu stricto and basophilous lichens, especially in the case of rare species. Of the species found in the study area, the following are definitely basophilous: Arthonia lapidicola, Arthopyrenia inconspicua, Candelariella aurella, Dermatocarpon miniatum, Lecanora dispersa, Lecidella stigmatea, Protoblastenia rupestris, Verrucaria muralis and V. nigrescens. These species grow not only on pure limestone, but also on other limy rocks or substrates covered with lime dust, as well as on silicaceous rocks containing calcium compounds (Wirth 1995). For example, Arthopyrenia inconspicua can grow also on basalts (see Nowak & Tobolewski 1975). Nitrophilous lichens require a large amount of nitrogen compounds for their life processes and are not strongly restricted to calcareous habitats. On the study area this group is represented mainly by species from genera Caloplaca (C. decipiens, C. saxicola), Phaeophyscia (Ph. nigricans, Ph. orbicularis), Physcia (Ph. adscendens, Ph. caesia) and Xanthoria (X. candelaria, X. elegans, X. parietina). All these species are widely distributed and abundant, especially in synanthropic habitats. 4.3. Epigeic (terricolous) lichens As it was explained in chapter 4.1, the present study included only those epigeic species that were connected with limestone outcrops and occured within investigated objects or in the immediate vicinity. The study therefore did not include such potential habitats for calciphilous and basophilous lichens as country roadsides or xerothermic grasslands on exposed slopes. In total, 33 epigeic species were found, of which most were very rare in the study area, occuring only at one or two localities. Half of these species were exclusively epigeic. This group included the following, together with the number of localities at which they were found: Agonimia gelatinosa (1), Bilimbia lobulata (1), Collema limosum (2), C. tenax (8), Endocarpon adscendens (1), E. pusillum (1), Fulgensia fulgens (1), Lempholemma chalazanum (4), L. polyanthes (1), Leptogium tenuissimum (2), Peltigera didactyla (1), P. rufescens (7), Placidium rufescens (1), P. squamulosum (1), Staurothele succedens (1), Thelocarpon impressellum (1) and Toninia sedifolia (2). One of them, Staurothele succedens, typically grows directly on rocks. In the study area, however, it was found exclusively on wet soil in a water seepage at the bottom of a quarry. The other 16 taxa are considered as passing species, which were also found growing on other substrates (Tab. 5). Of these, nine species were also epibryophytic. Most of them grew on both soil and moss at the same locality. 28 Maria-Kossowska 28 Table 5. Epigeic passing species growing on various types of substrates Species occuring on soil and mosses Cladonia glauca Cladonia pocillum Collema auriforme Lepraria eburnea Lepraria vouauxii Leptogium gelatinosum Leptogium lichenoides Leptogium schraderi Peltigera praetextata Species occuring on soil and rocks Aspicilia contorta Aspicilia moenium Collema crispum Lepraria eburnea Lepraria lobificans Lepraria vouauxii Protoblastenia rupestris In spite of their low participation in studied lichen flora, terricolous lichens deserve special attention. This group inlude a number of species very rare in Poland, and some of them had never before been found in Lower Silesia. They are: Agonimia gelatinosa, Endocarpon adscendens, E. pusillum, Fulgensia fulgens, Lempholemma polyanthes, Leptogium schraderi, Placidium rufescens and Thelocarpon impressellum. 4.4. Epibryophytic (muscicolous) lichens Epibryophytic lichens occupy a specific type of habitat. They are found mainly on patches of living moss that grow either on rock surfaces or on the soil. In some studies, this group is not treated separately, and the species in it are classified as either saxicolous or terricolous. In the present study, the group is treated separately in order to emphasize their undirect connexion with rock or soil. On the investigated area, epibryophytic species made the littest and the less specific group of lichens (16% of lichen flora). Exclusively epibryophytic lichens have still lower importance only six such species were found: Bacidia bagliettoana, Caloplaca cerina var. muscorum, Mycobilimbia pilularis, M. tetramera, Physconia grisea and Synalissa symphorea (less than 5% of lichen flora). The other species were also found growing on other types of substrates (Tab. 6). Of the exclusively epibryophytic species identified, those with the highest relative abundance were Bilimbia sabuletorum and Mycobilimbia tetramera. Both of these species were found growing on patches of epilithic and epigeic mosses at seven localities in the study area. The other exclusively epibryophytic species were each found at only one locality. This was also true for Caloplaca cerina var. muscorum, which is common in other calcareous regions in Poland (see Nowak 1961). 29 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 29 Table 6. Epibryophytic passing species growing on various types of substrates Species occuring on mosses and soil Cladonia glauca Cladonia pocillum Collema auriforme Lepraria eburnea Lepraria vouauxii Leptogium gelatinosum Leptogium lichenoides Leptogium schraderi Peltigera praetextata Species occuring on mosses and rocks Diploschistes muscorum Bilimbia sabuletorum Leptogium intermedium Lepraria eburnea Lepraria vouauxii Physcia adscendens Xanthoria candelaria Xanthoria parietina Among the muscicolous lichens, a number of gelatinous lichens containing a cyanobacterial symbiont, usually belonging to the genus Nostoc have been recorded. They are: Synalissa symphorea, Collema auriforme, Leptogium gelatinosum, L. intermedium, L. lichenoides and L. schraderi. All of these species are uncommon or rare in Poland. 4.5. Nitrophilous lichens General remarks. All living organisms require nitrogen to live. Only a few, however, are able to fix atmospheric elemental nitrogen. This ability is limited to several groups of bacteria, including cyanobacteria and actinomycetes. All other organisms have to depend on nitrogenous compounds as a source of nitrogen. Lichens that contain a nitrogen-fixing cyanobacterium (e.g. Nostoc) as a primary or secondary symbiont can grow in the absence of nitrogenous compounds. Other taxa require the presence of nitrogen in the substrate. However, excessive amounts of nitrogenous compounds can also limit the growth of many lichen species, as happens on rock surfaces covered with thick layers of bird droppings, or in areas that are encrusted with artificial fertilizers from nearby cultivated fields. These habitats are occupied by only a few lichen species that are extremely tolerant of high levels of nitrogenous compounds. Many of these species can grow even in the presence of uric acid, which is toxic for most other organisms. These species are able to transform uric acid into ammonium ions and carbon dioxide (Nash III 1996). They are usually referred to as nitrophilous species; Wirth (1980) called them extremely nitrophilous. 30 Maria-Kossowska 30 Table 7. Classification of lichens in terms of substrate eutrophy (Wirth 1980) Substrate Lichen not eutrophicated moderately eutrophicated rather eutrophicated strongly eutrophicated extremely eutrophicated anitrophilous moderately nitrophilous rather nitrophilous strongly nitrophilous extremely nitrophilous Although it is generally not difficult to distinguish extremely nitrophilous lichen species, the degree of nitrophilia in other species (or in what amounts they require nitrogenous compounds) is not so clear. There are differences in opinion among lichenologists about how the particular taxa should be classified. The most exhaustive classification of nitrophilous lichens was compiled by Wirth (1980, 1995). He distinguished five classes of nitrophilous lichens, depending on the trophic level of the habitats in which they occure (Tab. 7). However, the system is based on qualitative criteria, not quantitative ones. Subject interpretation therefore remains a problem. Participation of nitrophilous species in studied lichen flora. In spite of the reservations mentioned above, the degree of nitrophilia of the descriebed lichen flora was analysed in accordance with Wirths species diagnoses (op.cit.) and supplemented by informations published by Nimis (1993). The lichens found in the study area could be assigned to three groups (Tab. 8). The group I corresponds to the class of anitrophilous species in the system proposed by Wirth, the group II contains moderately nitrophilous and rather nitrophilous species and the group III consists of strongly nitrophilous and extremely nitrophilous species. Not all of the lichen taxa could be satisfactorily classified because their habitat preferences could not be accurately determined. When classified using the three groups listed above, the lichen flora of the study area appeared diverse in terms of the degree of nitrophilia. Among the reasons for this is that the localities examined in this study were themselves very diverse in terms of mineral content, and contained widely differing amounts of nitrogenous compounds. The objects with the highest levels of eutrophication were those immediately next to cultivated fields and stone quarries in which garbage was illegally dumped. Other habitats that was rich in nitrogenous compounds included the upper reaches of exposed rocks and the high walls of quarries. These sites were often covered by bird droppings. The lichen species dominating in these habitats belong to group III, including Caloplaca decipiens, Phaeophyscia orbicularis and Protoparmeliopsis muralis. Species belonging to group I made up the smallest part of the calcicolous lichen flora in the study area. Many of these species are rare in Poland, such 31 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 31 Table 8. Variation of degree of nitrophilia in calcicolous lichen flora Group Description Species (number of localities) I Species of habitats contained relatively low amounts of nitrogenous compounds Aspicilia calcarea (3), Clauzadea monticola (7), Collema callopismum (1), Endocarpon pusillum (1), Gyalecta jenensis (10), Hymenelia prevostii (2), Mycobilimbia pilularis (1), Mycobilimbia tetramera (7), Polyblastia albida (2), P. theleodes (1), Sarcogyne clavus (2), Thelidium papulare (2). II Species of habitats contained moderate amounts of nitrogenous compounds Acarospora cervina (6), A. glaucocarpa (5), A. macrospora (1), Aspicilia contorta subsp. contorta (19), A. c. subsp. hoffmanniana (5), Bilimbia sabuletorum (7), Caloplaca cerina var. cerina (1), C. chalybaea (3), C. cirrochroa (4), C. holocarpa (3), C. lactea (3), C. saxicola (1), C. teicholyta (2), C. variabilis (3), Candelariella aurella (19), C. vitellina (2), Catillaria chalybeia (6), C. lenticularis (12), Collema crispum (2), Dermatocarpon miniatum (3), Diplotomma alboatrum (3), Lecidella carpathica (5), L. stigmatea (23), Peltigera didactyla (1), Phaeophyscia sciastra (2), Physconia grisea (1), Protoblastenia rupestris (33), Rinodina bischoffii (3), Sarcopyrenia gibba (1 ), Synalissa symphorea (1), Verrucaria muralis (12), V. nigrescens (15). III Species of habitats rich in nitrogenous compounds Caloplaca citrina (15), C. decipiens (4), Lecania erysibe (1), Lecanora albescens (18), Phaeophyscia nigricans (5), Ph. orbicularis (8), Physcia adscendens (7), Ph. caesia (3), Protoparmeliopsis muralis (6), Xanthoria candelaria (1), X. elegans (6), X. parietina (3). as Collema callopismum, Endocarpon pusillum and Polyblastia theleodes. These species were found at only a small number of localities. This may be a result of the general eutrophication of natural environment due to human activity. 4.6. Classification of calcareous microhabitats Both types of analyzed objects, quarries and natural outcrops of calcareous rocks, varied widely in age, origin and ecological conditions. On each locality one could distinguish a range of microhabitats that complexly interacted with each other in terms of time and space. In quarries they include: rock faces (both exposed and sheltered), blocks at the base of the walls, small stones and rock debris, rock-waste and soil on the walls and the bottom of a quarry, which may be made of stones or covered by a layer of the soil. The microhabitats in natural outcrops also differed widely in terms of the availability of sunlight and water. Among others, the upper reaches of exposed rocks fertilized by birds and rocks sittuated in vicinity of cultivated fields could be distingiushed. These two microhabitats were particularly rich in nitrogen and phosphorus compaunds. 32 Maria-Kossowska 32 Table 9. Classification of the epilithic microhabitats I. Anthropogenic objects Class Microhabitat Species 1. Fresh rock faces (unweathered and) in quarries abandoned in the past few years. Aspicilia contorta subsp. contorta, A. moenium, Caloplaca citrina, C. holocarpa, Candelariella aurella, Catillaria lenticularis, Eiglera flavida, Lecanora albescens, L. dispersa, Lecidella stigmatea, Protoblastenia rupestris, Sarcogyne regularis. 2. ‘Mature’ rock faces (weathered and exposed to the sun) in quarries that have been abandoned for at least ten years, but not long enough for trees to grow. Acarospora cervina, A. glaucocarpa, Aspicilia contorta subsp. contorta, A. contorta subsp. hoffmanniana, Caloplaca citrina, C. crenulatella, C. velana, Candelariella aurella, Catillaria lenticularis, Clauzadea monticola, Dermatocarpon miniatum, Diploschistes gypsaceus, Farnoldia hypocrita, Hymenelia epulotica, Lecanora albescens, L. dispersa, Lecidella carpathica, L. stigmatea, Protoblastenia rupestris, Psorotichia schaereri, Rhizocarpon petraeum, Sarcogyne regularis, Thelidium cataractarum, Verrucaria hochstetteri, V. deminuta, V. muralis, V. nigrescens, V. nigroumbrina, V. obfuscans. 3. Upper reaches of ‘mature’ rock faces frequently visited by birds. as in I.2., together with: Caloplaca decipiens, Diplotomma alboatrum, Lecidella carpathica, Phaeophyscia nigricans, Ph. orbicularis, Physcia caesia, Protoparmeliopsis muralis, Sarcopyrenia gibba, Xanthomendoza fallax, Xanthoria elegans. 4. Shaded rock faces in abandoned quarries of several score years, possibly overgrown with trees, and north-facing rocks in younger objects. Aspicilia contorta subsp. contorta, Caloplaca citrina, Collema fuscovirens, Gyalecta jenensis, Lepraria sp. div., Polyblastia theleodes, Protoblastenia rupestris, Sarcogyne clavus, Thermutis velutina, Verrucaria dolosa, V. hochstetteri, V. viridula. 5. Stone blocks exposed at the base of the walls of the quarry. Acarospora glaucocarpa, Aspicilia contorta subsp. contorta, A. moenium, Caloplaca citrina, C. crenulatella, Candelariella aurella, Catillaria lenticularis, Lecanora albescens, L. dispersa, Lecidella stigmatea, Phaeophyscia nigricans, Ph. orbicularis, Protoblastenia rupestris, Protoparmeliopsis muralis, Sarcogyne regularis, Verrucaria deminuta, V. muralis, V. nigrescens, Xanthoria elegans, X. parietina. 6. Little stones. Acarospora glaucocarpa, Aspicilia moenium, Candelariella aurella, Eiglera flavida, Lecanora dispersa, Lecidella stigmatea, Protoblastenia rupestris, Sarcogyne regularis, Staurothele hymenogonia, Verrucaria muralis, V. nigrescens. 33 II. Natural outcrops Class Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains Microhabitat 33 Species 1. Insolated rocks. Acarospora macrospora, Aspicilia calcarea, A. coronata, A .contorta subsp. hoffmanniana, Diplotomma alboatrum, Caloplaca chalybaea, C. velana, Candelariella aurella, Catillaria lenticularis, Clauzadea monticola, Collema fuscovirens, Dermatocarpon miniatum, Lecanora crenulata, L. dispersa, Lecidella stigmatea, Placynthium nigrum, P. tremniacum, Protoblastenia incrustans, P. rupestris, Rinodina bischoffii, Sarcogyne regularis, Verrucaria nigrescens, V. obfuscans. 2. Top parts of insolated rocks, frequently visited by birds. as in II.1., together with: Caloplaca coronata, C. decipiens, Phaeophyscia nigricans, Ph. orbicularis, Protoparmeliopsis muralis, Rinodina teichophila 3. Exposed rock outcrops near cultivated fields in which nitrogen-based and phosphorusbased artificial fertilizers are used. Aspicilia contorta subsp. contorta, Caloplaca decipiens, C. teicholyta, C. velana, Candelariella aurella, C. vitellina, Catillaria lenticularis, Lecanora albescens, L. campestris, L. dispersa, Lecidella carpathica, L. stigmatea, Phaeophyscia orbicularis, Ph. sciastra, Physcia caesia, Ph. adscendens, Protoblastenia rupestris, Protoparmeliopsis muralis, Sarcogyne regularis, Scoliciosporum umbrinum, Verrucaria procopii, Xanthomendoza fallax, Xanthoria candelaria, X. elegans, X. parietina. 4. Shaded rock outcrops in wooded areas. Arthopyrenia inconspicua, Caloplaca cirrochroa, Collema crispum, Dermatocarpon miniatum, Gyalecta jenensis, Hymenelia prevostii, Lemmpopsis arnoldiana, Lepraria sp. div., Leptogium intermedium, L. plicatile, Polyblastia albida, Protoblastenia rupestris, Sarcogyne regularis, S. clavus, Thelidium incavtum, Verrucaria hochstetteri, V. dolosa. Particular microhabitats differ in terms of the species number and the characteristic composition of the lichen flora. Some species are found in all of the microhabitats identified (e.g. Lecanora dispersa and Protoblastenia rupestris), other species prefer a single microhabitat or a small number of similar habitats (e.g. Gyalecta jenensis growing only in shaded places). The temporary classification of the epilithic microhabitats identified in the study area is presented in Table 9. They are divided into two classes in therms of their natural or anthropogenic origine. Anologous microhabitats at anthropogenic and natural localities (e.g. shady natural rocks and sheltered walls in 34 Maria-Kossowska 34 quarries) differed from each other, especially in terms of the species number. Microhabitats at natural localities were generally characterized by a higher number of species. At anthropogenic localities, on the other hand, the number of more or less nitrophilous species was higher. This is typical for habitats that have been created or altered by human activity. The lichen flora found at anthropogenic localities corresponds to that found on artificial substrates such as concrete. At natural localities, this was also somewhat true only for rocks sittuated near cultivated fields. The epigeic and epibryophytic habitats identified in the study area form the separate group. As with the epilithic microhabitats, this group is not homogenous and particular epigeic and epibryophytic microhabitats differed among themselves. There are five microhabitats distinguished, each characterized by a different combination of species (Tab. 10). Because most epigeic and epibryophytic lichens were found at only one or a few localities, it was impossible to construct a more detailed classification of the microhabitats in which they occurred. Table 10. Distinguished epigeic microhabitats Microhabitat Species 1. Rock waste and soil on exposed walls of quarries or in crevices of natural outcrops. Aspicilia contorta, Endocarpon adscendens, Lempholemma chalazanum, L. polyanthes, Leptogium gelatinosum, L. tenuissimum, Protoblastenia rupestris. 2. Bare soil in open places or in patches of short, xerothermic grassland. Collema limosum, C. tenax, Endocarpon pusillum, Fulgensia fulgens, Leptogium tenuissimum, Placidium rufescens, P. squamulosum, Toninia sedifolia. 3. Moss-covered soil in exposed and relatively dry places. Aspicilia moenium, Cladonia glauca, C. pocillum, Leptogium lichenoides, L. schraderi, Peltigera rufescens. 4. Moss-covered soil in shaded places. Leptogium gelatinosum, Peltigera praetextata. 5. Humus soil among mosses in relative moist places. Agonimia gelatinosa, Staurothele succedens, Thelocarpon impressellum. 5. Comparison of the calcicolous lichen flora in the Western and Eastern Sudety Mountains Although habitat conditions in the calcareous areas of the Western and Eastern Sudety Mts were largely similar, there were significant differences between these regions in both the present-day lichen flora and the total lichen flora including historical data (Fig. 6). The Western and Eastern Sudetes differed in terms of 35 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 35 B A 0 20 40 60 1 80 100 2 120 140 160 180 3 Fig. 6. Comparison of the calcicolous lichen flora in the Western and Eastern Sudety Mts A total lichen flora; B present-day lichen flora; 1 species found in both areas; 2 species exclusive for the Western Sudetes; 3 species exclusive for the Eastern Sudetes both number of lichen species and species composition. Of the 129 species identified in this study, only 54 were found in both areas. The other 75 are presently known from only one of the regions at present (although some of them had been widely distributed in the past, as for example Acarospora macrospora, Caloplaca variabilis, Collema crispum, Polyblastia albida, Thelidium incavatum, Toninia sedifolia and Verrucaria foveolata). Many of the taxa found in both areas were species with broad habitat preferences that are common in all calcareous regions in Poland as well as on artificial calcareous substrates, such as: Aspicilia contorta subsp. contorta, Caloplaca citrina, Candelariella aurella, Lecanora albescens, L. crenulata, L. dispersa, Lecidella stigmatea, Phaeophyscia orbicularis, Physcia adscendens, Protoblastenia rupestris, Sarcogyne regularis, Verrucaria muralis, V. nigrescens and Xanthoria parietina, as well as typical calciphilous species that are commonly found on calcareous rock formations in other areas in Poland, such as Acarospora cervina, A. glaucocarpa, Caloplaca velana, Collema fuscovirens, Gyalecta jenensis and Rinodina bischoffii. Several species, however, are characteristic for the Sudety Mountains and are rarely found in other regions of Poland. These include Collema auriforme, Eiglera flavida, Leptogium gelatinosum, L. plicatile and Thelidium gisleri. 36 Maria-Kossowska 36 In the calcareous area of the Western Sudety Mountains, a total of 111 calcicolous lichen species were found, of which 57 were not found in the Eastern Sudety Mountains. Most of the rare and endangered species mentioned in this study occured just in this area. These include Agonimia gelatinosa, Arthopyrenia inconspicua, Endocarpon adscendens, Fulgensia fulgens, Lemmopsis arnoldiana, Lempholemma polyanthes, Leptogium intermedium, Placidium rufescens, Polyblastia theleodes, Sarcopyrenia gibba, Staurothele succedens and Thermutis velutina. In the Eastern Sudety Mountains, the present-day lichen flora is ditinctly poorer than in the Western Sudetes. A total of 72 calcicolous lichen species were identified during the study, of which only 18 were not found in the Western Sudety Mts. Some of these were also rare in Poland, but they made up only a small part of the local lichen flora. These include Collema callopismum, Endocarpon pusillum, Leptogium schraderi, Staurothele hymenogonia, Synalissa symphorea, Thelidium cataractarum and T. incavatum. The Eastern Sudety Mts differed also in small number of epigeic species, especially those that prefer xerothermic grassland habitats. Only four such species were found here: Cladonia pocillum, Collema tenax, Endocarpon pusillum and Leptogium schraderi. In the Western Sudetes, eight epigeic species occured: Cladonia pocillum, Collema tenax, Endocarpon adscendens, Fulgensia fulgens, Leptogium tenuissimum, Placidium rufescens, P. squamulosum and Toninia sedifolia. The significant richness and high diversity of calcicolous lichen flora of the Western Sudety Mts can be explained as a result of lower metamorphism of the Cambrian limestones (see chapter 1.3). Owing to this, they are less compact than those in the Eastern Sudetes, so the calcium carbonate that they contain is more easily available. Another reason is that there are more exposed natural calcareous rock formations in the Western Sudety Mts. These sites served as reservoirs of epilithic lichen species which could colonize abandoned quarries. 6. Threat to calcicolous lichen flora in the Sudety Mountains 6.1. Factors of threat Epilithic lichen species are generally considered to be in less danger of extinction than species which grow on trees, mainly because they are more resistant to air pollution. Among epilithic species, calcicolous species are considered to be the most resistant to acid rain and the sulfuric acid it contains because of the buffering capacity of calcareous substrates (Fa³tynowicz 1992). Nevertheless, the calcicolous lichen flora of the Sudety Mountains is severely threatened due to a number of other direct and indirect factors. As it was mentio- 37 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 37 ned above, the majority of species identified in this study were found at only one or two localities (category very rare in the species frequency scale; see chapter 2). Only few of the species were frequent in the study area, and these were mainly species with broad habitat preferences. No species found could be classified as common according to the species category scale (occuring at over 75% of the investigated objects). Among the factors that threaten the calcicolous lichen flora of the Sudety Mounts are the following: - anthropogenic and natural outcrops of calcareous rock are sparsely scattered in throughout the region, so particular populations of calcicolous lichen species are isolated; - plant succession in abandoned quarries means that exposed rock surfaces are increasingly shaded, which is unfavorable for the photophilous species that make up the greater part of the calcicolous lichen flora; - intensified exploitation of active quarries and re-activation of abandoned quarries it is a particular problem because there are so few exposed natural calcareous rock outcrops in the region and many of the most valuable species are found in or near abandoned quarries (including Fulgensia fulgens, which has been found at only few other sites in Poland); - illegal dumping of garbage in abandoned quarries causes eutrophication of epilithic and epigeic habitats, which is responsible for the disappearance of species that cannot tolerate high concentrations of nitrogenous compounds in the substrate; - intensified agricultural activity in the region and the greater use of artificial fertilizers it entails are especially unfavorable for lichen species growing on exposed field boulders or in xerothermic grasslands near cultivated fields; - air pollution is probably also unfavorable for calcicolous lichen species, though not as much as it is for lichen species that grow on trees. Only the first two have natural character; other factors listed above are connected with human pressure on the natural environment, the effects of which have been increasing over the past few decades. 6.2. Participation of rare and endangered species Of the 174 calcicolous lichen species found in the Sudety Mountains till present, 84 species are either extinct or threatened in Poland and are entered to the Polish red list of lichens (Cieliñski et al. 2003; Tab. 11). Among them there are five species that are believed to be extinct in the country (category RE) and numerous group of endangered species (categories CR and EN, together 26 species). Additionaly, species listed under category DD (data defi- 38 Maria-Kossowska 38 cient) deserve special attention, because their status of threat in Poland is probably very high. Table 11. Calcicolous lichen species found in the Sudety Mts that are entered in the Polish red list of lichens (Cieliñski et al. 2003) Category of threat Species RE Staurothele guestphalica, Thelidium aphanes, Verrucaria cincta, V. longicollis, V. saprophila CR Bacidina egenula, Endocarpon adscendens, Fulgensia fulgens, Megaspora verrucosa, Opegrapha calcarea Pseudosagedia byssophila, Rhizocarpon umbilicatum, Squamarina cartilaginea, Staurothele succedens, Toninia aromatica EN Arthopyrenia inconspicua, Catapyrenium daedaleum, Collema cristatum, C. callopismum, C. polycarpon, Heppia lutosa, Hymenelia epulotica, Lemmopsis arnoldiana, Leptogium subtile, Peccania coralloides, Placidium rufescens, Polyblastia sepulta, Sarcogyne clavus, Squamarina lentigera, Synalissa symphorea, Toninia candida VU Acarospora cervina, Bacidia trachona, Caloplaca cerina var. cerina, C. variabilis, Catapyrenium cinereum, Catillaria chalybeia, Dermatocarpon miniatum, Diploschistes gypsaceus, Diplotomma alboatrum, Endocarpon pusillum, Lempholemma polyanthes, Leptogium gelatinosum, L. schraderi, Mycobilimbia pilularis, Opegrapha dolomitica, Peltigera praetextata, Polyblastia theleodes, Protoblastenia incrustans, Rhizocarpon petraeum, Solorina saccata, Staurothele hymenogonia, Thelidium decipiens, Thelocarpon impressellum, Verrucaria viridula, Xanthomendoza fallax, Xanthoria calcicola NT Acarospora macrospora, Arthonia lapidicola, Bilimbia lobulata, Caloplaca cirrochroa, Catillaria lenticularis, Collema auriforme, Eiglera flavida, Gyalecta jenensis, Lempholemma chalazanum, Leptogium tenuissimum, Placidium squamulosum, Placynthium nigrum, Psorotichia schaereri, Thelidium incavatum, Th. papulare, Toninia sedifolia LC Acarospora glaucocarpa, Agonimia gelatinosa, Leptogium lichenoides, L. plicatile, Phaeophyscia sciastra DD Acrocordia conoidea, Aspicilia farinosa, Mycobilimbia lurida, Porocyphus rehmicus, Thermutis velutina, Verrucaria murina Species in bold print were found in the region during the study. The high number of taxa that are considered as endangered in a supra-regional scale, such as sensitive, stenotopic and generally rare species, speaks well for environmental values of given region. However, in the case of calcicolous lichen flora of the Sudety Mts the time factor is very important. Most of the species belonging to categories CR and EN were reported only in historical surveys are no longer found in the area. Furthermore, when found, they occurred at only one or two localities, where they made up only a tiny part of 39 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 39 the lichen vegetation. It seems therefore only a matter of time before they, too, become extinct. Of the 129 calcicolous lichen species identified in the present study, about 40% are threatened or endangered in the supra-regional scale. Among them, lichens of two categories: Vulnerable (VU 20 taxa) and Near Threatened (NT 16 taxa) dominate. A comparison of the relative abundance of threatened and endangered species in the total and present-day lichen floras preveals that the most valuable species are being lost (Fig. 7). B A 0 20 1 40 60 2 3 80 100 4 120 5 140 6 160 7 180 8 Fig. 7. Number of lichen species in particular Red List categories (Cieliñski et al. 2003) A total lichen flora; B present-day lichen flora 1 Regionally Extinct (RE); 2 Critically Endangered (CR); 3 Endangered (EN); 4 Vulnerable (VU); 5 Near Threatened (NT); 6 Least Concern (LC); 7 Data Deficient (DD); 8 not threatened. 7. Summary and conclusions 1. The aim of the present study was to determine the current state of the lichen flora inhabiting crystalline limestone substrates in the Sudety Mountains. The investigations vere made in the two most extensive calcareous areas in Lower Silesia: the nie¿nik Metamorphic Region in the Eastern Sudetes and the Góry Kaczawskie Mts together with the adjacent portions of the Pogórze Kaczawskie Foothills and Pogórze Izerskie Foothills in the Western Sudetes. The objects investigated in these areas included both natural outcrops of calcar- 40 Maria-Kossowska 40 eous rock as well as limestone quarries. In all, 45 localities were examined 27 localities in the Western Sudety Mts and 18 in the Eastern Sudety Mts. 2. In this and previous surveys, 174 calcicolous lichen species have been found in the study areas, of which 129 are represented in the present-day lichen flora. The species examined included epilithic lichens as well as epigeic and epibryophytic species growing on or near exposed calcareous rocks. Many of these species are rare in Poland. Some had never before been found in the Sudety Mountains or the province of Lower Silesia. These include Agonimia gelatinosa, Endocarpon adscendens, Fulgensia fulgens, Lemmopsis arnoldiana, Lempholemma polyanthes, Leptogium intermedium, Placidium rufescens, Sarcopyrenia gibba, Staurothele succedens and Thelocarpon impressellum. 3. The present-day lichen flora includes species mentioned in the historical literature and confirmed during the study, as well as 53 new species. Most of them had never been found in the Sudety Mts but a few had been found previously in this region, but only on different types of substrate. These species can therefore be considered new for limestones of the Sudety Mts. 4. 45 species reported from the study area in 19th and the early half of 20th centuries were not re-discovered. They might disapear due to physical destruction of the former locality, for example reactivation of the formerly abandoned quarry. Another reason is that rock surfaces that had been once exposed were now overgrown with trees, which limits the availability of sunlight. Some of these species, however, may not be entirely extinct in the study areas, and may be found at other localities in future surveys. 5. The small group of widespread and common species that can colonize a wide range of natural and artificial calcareous substrates is the essential part of investigated lichen fora. They include: Aspicilia contorta subsp. contorta, Caloplaca citrina, Candelariella aurella, Lecanora albescens, Lecanora dispersa, Lecidella stigmatea, Sarcogyne regularis, Protoblastenia rupestris, and Verrucaria nigrescens. Each of these species was found at at least 15 of the 45 localities examined. 6. About half of the lichen species are very rare in the Sudety Mts, and were found at only one or two sites in the study area. Some of them are rare in Poland as a whole, such as Agonimia gelatinosa, Arthopyrenia inconspicua, Caloplaca crenulatella, Endocarpon adscendens, E. pusillum, Fulgensia fulgens, Lemmopsis arnoldiana, Leptogium intermedium, L. plicatile, L. tenuissimum, Placidium rufescens, Sarcopyrenia gibba, Thelidium gisleri, Toninia sedifolia. Some of these species had not been reported from this area till now. 7. The lichens identified in the study area could be assigned to three groups depending on the level of nitrogenous compounds in the substrate on which they were found: 41 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 41 - species that prefer habitats with a low level of nitrogenous compounds; - species typical for habitats moderately rich in nitrogenous compounds; - species of habitats extremly rich in nitrogen. Taxa belonging to the first group made up the smallest part of the calcicolous lichen flora in the study area. This may be a result of the general eutrophication of natural environment due to human activity. 8. Within the natural and anthropogenic objects included in this study a range of microhabitats was distinguished that differed in terms of the species number and the characteristic composition of the lichen flora. They include: rock faces in quarries (both exposed and sheltered), blocks at the base of the walls, small stones and rock debris, rock-waste and soil on the walls and the bottom of a quarry, natural outcrops (also exposed and sheltered) etc. The microhabitats are divided into two classes in therms of their natural or anthropogenic origine. 9. The calcicolous lichen flora of the Western Sudety Mts was significantly richer and more diverse than that of the Eastern Sudety Mts and corresponds with other calcareous regions in Poland (e.g. Pieniny Mts, Wy¿yna KrakowskoWieluñska Upland and Góry wiêtokrzyskie Mts). This is probably caused by lower metamorphism (epimetamorphism) of the limestones occuring here. Owing to this, they are less compact than marbles in the Eastern Sudety Mts and the calcium carbonate is more easily available. 10. The lichen fora of investigated calcareous areas in the Sudety Mts is very interesting, rich and diverse. However, the calcicolous lichens occuring here are highly threatened and many species were found at only one or two localities. Factors that pose an indirect threat include air pollution, to which calcicolous lichen species are also sensitive. Other factors pose a direct threat, such as destruction of the localities, for example the activation of abandoned quarry sites. Many of the quarries examined in the present study may be reactivated in order to satisfy increasing demand, or to provide employment in times of economic crisis. Those sites that are most valuable from a lichenological point of view should therefore be protected as reserves or documentation stands. 8. References B ¥C -B RONOWICZ J. 1997. Opady atmosferyczne. In: W. P AWLAK (ed.), Atlas l¹ska Dolnego i Opolskiego. Wroc³aw University, Wroc³aw. C IELIÑSKI S., C ZY¯EWSKA K. 1992. Problemy zagro¿enia porostów w Polsce. Wiad. Bot. 36(12): 517. C IELIÑSKI S., C ZY¯EWSKA K., F ABISZEWSKI J. 2003. Czerwona lista porostów wymar³ych i zagro¿onych w Polsce. In: K. C ZY¯EWSKA (ed.). Zagro¿enie porostów w Polsce. Monogr. Bot. 91: 1449. C IELIÑSKI S., F A£TYNOWICZ W. (eds) 1993. Atlas of geographical distribution of lichens in Poland. Part. I. W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków, 67 pp. 42 Maria-Kossowska 42 C ZY¯EWSKA K. 1986. Flora porostów naziemnych w Za³êczañskim Parku Krajobrazowym (Wy¿yna Wieluñska). Acta Univ. Lodz., Folia Sozol. 2: 315 341. D EGELIUS G. 1954. The lichen genus Collema in Europe. Symb. Bot. 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Zarys klimatu Polski. Bogucki Wyd. Naukowe, Poznañ, 302 pp. WÓJCIAK H. 1987. Flora porostów wapiennych zboczy w Dobrem ko³o Kazimierza Dolnego nad Wis³¹. Annales UMCS, sec. C, 42(9): 103108. ZAJ¥C A. 1978. Za³o¿enia metodyczne Atlasu rozmieszczenia rolin naczyniowych w Polsce. Wiad. Bot. 22(3): 145155. 45 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 45 9. Streszczenie Porosty ska³ wapiennych w polskiej czêci Sudetów 1. Badania objête niniejszym opracowaniem podjête zosta³y w celu poznania aktualnego stanu lichenobioty zwi¹zanej z wyst¹pieniami wapieni krystalicznych w Sudetach. Przeprowadzono je na dwóch najwiêkszych obszarach wapiennych w Sudetach w obrêbie tzw. Metamorfiku nie¿nika (Góry Bialskie, Góry Z³ote i Masyw nie¿nika ³¹cznie z pasmem Krowiarek) w Sudetach Wschodnich oraz w Górach Kaczawskich (wraz z bezporednio przyleg³ymi fragmentami Pogórza Izerskiego i Pogórza Kaczawskiego) w Sudetach Zachodnich; ich teren ograniczony by³ bezporednio wystêpowaniem naturalnych (ska³ki) lub sztucznych (kamienio³omy) wychodni ska³ wapiennych. Przeanalizowano ³¹cznie 45 obiektów 27 w czêci zachodniej i 18 w czêci wschodniej Sudetów. 2. Przeprowadzona analiza lichenobioty nawapiennej w Sudetach opiera siê g³ównie na materiale zebranym w terenie w latach 19961998. Dane historyczne pochodz¹ z opracowañ Flotowa (1850, 1851), Körbera (1855), Steina (1879, 1889), Eitnera (1896, 1901, 1911) i Fabiszewskiego (1968). Wykorzystano te¿ okazy znajduj¹ce siê w herbarium Uniwersytetu Wroc³awskiego (WRSL) i zielniku Uniwersytetu Adama Mickiewicza (POZ). 3. Na obu badanych obszarach stwierdzono ogó³em 174 taksony porostów (Tab. 3, strona 17). Na liczbê tê sk³adaj¹ siê dwie podstawowe kategorie: lichenobiota historyczna porosty stwierdzone do roku 1970, podawane w literaturze lub zachowane w zbiorach zielnikowych. Liczy ³¹cznie 131 gatunków, w tym 108 gatunków pewnych (podanych z konkretnych stanowisk na badanym terenie) oraz 13 gatunków prawdopodobnych (wymienianych ogólnie jako rozpowszechnione lub czêste na obszarze l¹ska, lecz bez podania konkretnych stanowisk; do grupy tej wliczono wy³¹cznie gatunki znane wspó³czenie z terenu badañ). lichenobiota wspó³czesna, czyli porosty odnalezione w trakcie badañ terenowych w latach 19961998. Liczy 129 ³¹cznie gatunków, w tym taksony historyczne pewne potwierdzone wspó³czenie (63 gatunki), historyczne prawdopodobne (13 gatunków) oraz taksony nowe (53 gatunki). Tê ostatni¹ grupê tworz¹ zarówno porosty dot¹d w Sudetach nienotowane, jak i znane z tego obszaru, ale znajdowane dotychczas wy³¹cznie na ska³ach krzemianowych (gatunki nowe dla sudeckich wapieni). 46 Maria-Kossowska 46 4. Sporód 108 taksonów znanych z literatury 45 nie uda³o siê odnaleæ wspó³czenie. Ich zanik mo¿e byæ wywo³any likwidacj¹ stanowisk, np. przez uruchomienie kamienio³omu, lub te¿ diametraln¹ zmian¹ warunków siedliskowych (np. zacienienie ods³oniêtych niegdy powierzchni skalnych przez las). Pewn¹ rolê w zanikaniu poszczególnych gatunków prawdopodobnie wywar³a te¿ szeroko pojêta antropopresja, a zw³aszcza zanieczyszczenie powietrza, które oddzia³uje negatywnie tak¿e na porosty nawapienne. Byæ mo¿e tak¿e czêæ gatunków uznanych za wy³¹cznie historyczne zosta³a przeoczona w trakcie badañ terenowych, z uwagi na swe bardzo niepozorne rozmiary (np. gatunki z rodzaju Thelidium i Verrucaria), a dalsze, szczegó³owe poszukiwania doprowadz¹ do ich odnalezienia. 5. Trzon lichenobioty Sudetów stanowi niewielka grupa porostów pospolitych w na terenie ca³ego kraju, o szerokiej amplitudzie ekologicznej, zasiedlaj¹cych zarówno pod³o¿a naturalne, jak i antropogeniczne (beton) i czêsto uwa¿anych za pionierskie: Caloplaca citrina, Candelariella aurella, Lecanora dispersa, Lecidella stigmatea, Protoblastenia rupestris, Sarcogyne regularis, Verrucaria nigrescens i kilka innych. Gatunki te znane s¹ z co najmniej jednej trzeciej wszystkich badanych obiektów. Listê znacznie wzbogacaj¹ gatunki bardzo rzadkie w Sudetach, znane tu z 12 stanowisk. Grupa ta liczy oko³o 50% wszystkich porostów. S¹ wród nich taksony rzadkie w Polsce, np. Agonimia gelatinosa, Arthopyrenia inconspicua, Caloplaca crenulatella, Endocarpon adscendens, E. pusillum, Fulgensia fulgens, Lemmopsis arnoldiana, Leptogium intermedium, L. plicatile, L. tenuissimum, Placidium rufescens, Sarcopyrenia gibba, Thelidium gisleri, Toninia sedifolia i in. Czêæ z nich nie by³a dot¹d notowana na badanym obszarze. Przedstawiony obraz lichenobioty naskalnej, z nieliczn¹ grup¹ gatunków bardzo czêstych i czêstych oraz dominuj¹c¹ grup¹ gatunków bardzo rzadkich, jest typowy dla obszarów o mocno zró¿nicowanych warunkach ekologicznych i du¿ym rozproszeniu potencjalnych siedlisk, pomiêdzy którymi wystêpuj¹ rozleg³e nieci¹g³oci. Mo¿e jednak te¿ wiadczyæ o intensywnym procesie wymierania gatunków na badanym obszarze. 6. Dokonana analiza form morfologicznych wród porostów naskalnych badanego obszaru pozwoli³a na wyodrêbnienie szeciu form morfologicznych: - porosty skorupiaste epilityczne (cep) 63 gatunki, - porosty skorupiaste endolityczne (cen) i z endolitycznym podpleszem (cep/n) 12 gatunków, - porosty plakodiowe (plac) 4 gatunki, - porosty ³useczkowate (sq) 2 gatunki, 47 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 47 - porosty listkowate (fol) 14 gatunków, z czego jedn¹ trzeci¹ stanowi¹ porosty sinicowe z rodziny Collemataceae, - porosty nitkowate (fil) 1 gatunek. Sporód wymienionych, na szczególn¹ uwagê zas³uguje grupa porostów endolitycznych i z endolitycznym podpleszem. Grupa ta z regu³y jest szczególnie licznie reprezentowana na ska³ach wapiennych, jednak w Sudetach udzia³ w lichenobiocie nie jest znaczny (17,7%), a po odrzuceniu gatunków wytwarzaj¹cych wyran¹ plechê zewnêtrzn¹ i tylko endolityczne podplesze staje siê on wrêcz znikomy. Jest to cecha charakterystyczna dla Sudetów, wyró¿niaj¹ca j¹ sporód innych obszarów wapiennych w Polsce i zwi¹zana ze specyficznymi w³aciwociami pod³o¿a skalnego. Proces metamorfizacji, jakiemu uleg³y pierwotne ska³y osadowe, przekszta³caj¹c siê w wapienie krystaliczne, spowodowa³ ich du¿¹ zwiêz³oæ i twardoæ oraz relatywn¹ odpornoæ kryszta³ów wêglanu wapnia na rozpuszczanie. Takie pod³o¿e staje siê za bardzo trudne do kolonizacji dla gatunków endolitycznych. Podobne obserwacje poczyni³ Fröberg (1989) na twardych wapieniach ordowickich w po³udniowej Skandynawii. 7. Badan¹ lichenobiotê przeanalizowano pod wzglêdem tolerancji wobec obecnoci zwi¹zków azotu w pod³o¿u. Wydzielono trzy grupy siedliskowe porostów (Tab. 8, strona 31): - gatunki siedlisk nieznacznie wzbogaconych w zwi¹zki azotu, - gatunki siedlisk zawieraj¹cych umiarkowane iloci zwi¹zków azotu, - gatunki siedlisk bardzo bogatych w azot. Badane stanowiska s¹ niejednolite pod wzglêdem zawartoci zwi¹zków azotu. Najbardziej eutroficzne s¹ obiekty znajduj¹ce siê w bezporednim s¹siedztwie pól uprawnych oraz kamienio³omy, w których urz¹dzono dzikie wysypiska mieci. Zwiêkszon¹ koncentracj¹ zwi¹zków azotu, pochodz¹cych z odchodów ptasich, cechuj¹ siê równie¿ wierzcho³ki nas³onecznionych ska³ek i wysokich cian kamienio³omów. Znajduje to odbicie w dominacji na tych stanowiskach gatunków z grupy trzeciej, np.: Caloplaca decipiens, Phaeophyscia orbicularis, Protoparmeliopsis muralis i in. Najmniejszy udzia³ w lichenobiocie wspó³czesnej badanego obszaru maj¹ porosty z grupy pierwszej, np.: Aspicilia calcarea, Collema callopismum, Endocarpon pusillum, Hymenelia prevostii, Mycobilimbia pilularis, Polyblastia albida, P. theleodes, Thelidium papulare, co t³umaczyæ mo¿na ogóln¹ eutrofizacj¹ rodowiska przyrodniczego, wywo³an¹ szeroko pojêt¹ dzia³alnoci¹ cz³owieka. 8. Analizowane obiekty kamienio³omy i naturalne wychodnie ska³ wapiennych nie s¹ jednolite, zarówno pod wzglêdem wieku, pochodzenia, jak i panuj¹cych w nich warunków ekologicznych. W ich obrêbie wydzielono szereg mikrosiedlisk, wzajemnie siê zazêbiaj¹cych i pozostaj¹cych 48 Maria-Kossowska 48 w skomplikowanych relacjach czasowo-przestrzennych: zacienione i nas³onecznione ciany skalne w kamienio³omach, bloki skalne u podnó¿a cian, rumosz skalny, kamieniste lub pokryte gleb¹ dno kamienio³omu, naturalne ska³ki (równie¿ zró¿nicowane na nas³onecznione i zacienione) itp. Poszczególne mikrosiedliska ró¿ni¹ siê miêdzy sob¹ liczb¹ i charakterystyczn¹ kombinacj¹ zasiedlaj¹cych je porostów. Wród nich wyró¿niæ mo¿na gatunki ubikwistyczne obecne na wszystkich typach mikrosiedlisk (np. Lecanora dispersa i Protoblastenia rupestris), oraz gatunki przywi¹zane do jednego lub kilku pokrewnych ich typów (np. Gyalecta jenensis, wystêpuj¹ca wy³¹cznie w miejscach cienistych). Ze wzglêdu na pochodzenie wyró¿nione mikrosiedliska podzielono na dwie klasy: klasa I zwi¹zana jest z kamienio³omami, klasa II z naturalnymi wychodniami skalnymi (Tab. 9, strona 32). Analogiczne mikrosiedliska w obu klasach (np. zacienione naturalne ska³ki i zacienione ciany kamienio³omów) ró¿ni¹ siê nieco miêdzy sob¹, zw³aszcza pod wzglêdem liczby zasiedlaj¹cych je gatunków, przy czym z regu³y wiêcej porostów notowano na mikrosiedliskach naturalnych. W obiektach antropogenicznych wystêpuje za to wiêcej gatunków wymagaj¹cych do swego rozwoju obecnoci zwi¹zków azotu (porosty mniej lub bardziej nitrofilne). Ich zwiêkszony udzia³ typowy jest dla siedlisk utworzonych lub przekszta³conych przez cz³owieka i zbli¿a omawiane siedliska do sztucznych pod³o¿y wapiennych, np. betonu. Na stanowiskach naturalnych podobny sk³ad gatunkowy porostów maj¹ jedynie ska³y znajduj¹ce siê wród pól uprawnych. 9. Pomimo du¿ego podobieñstwa warunków siedliskowych panuj¹cych na obu badanych obszarach, pomiêdzy lichenobiot¹ nawapienn¹ w zachodniej i wschodniej czêci Sudetów istniej¹ doæ znaczne ró¿nice. Przejawiaj¹ siê one zarówno w ogólnej liczbie stwierdzonych taksonów, jak i sk³adzie gatunkowym na 129 odnalezionych wspó³czenie gatunków porostów, tylko 54 wspólne s¹ dla obu obszarów; pozosta³e 75 to taksony znane obecnie wy³¹cznie z jednego z nich. Wród gatunków wspólnych znajduj¹ siê przede wszystkim porosty o szerokiej amplitudzie ekologicznej, czêste na wszystkich obszarach wapiennych w Polsce i chêtnie przechodz¹ce na sztuczne pod³o¿a wêglanowe: Aspicilia contorta subsp. contorta, Caloplaca citrina, Candelariella aurella, Lecanora albescens, L. crenulata, L. dispersa, Lecidella stigmatea, Phaeophyscia orbicularis, Physcia adscendens, Protoblastenia rupestris, Sarcogyne regularis, Verrucaria muralis, V. nigrescens i Xanthoria parietina, oraz typowe gatunki kalcyfilne, równie¿ pospolicie wystepuj¹ce na pozosta³ych obszarach zalegania ska³ wêglanowych, np.: Acarospora cervina, A. glaucocarpa, Caloplaca velana, Collema fuscovirens, Gyalecta jenensis, Rinodina bischoffii i in. Na obszarze wystêpowania ska³ wapiennych w Sudetach Zachodnich stwierdzono ogó³em 111 gatunków porostów, z czego a¿ 57 to taksony wy³¹czne. 49 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 49 Lichenobiota nawapienna Sudetów Wschodnich jest znacznie ubo¿sza zanotowano tu wspó³czenie 72 gatunki porostów, w tym zaledwie 18 gatunków wy³¹cznych. Znacznie wiêksze bogactwo i zró¿nicowanie bioty porostów nawapiennych w G. Kaczawskich jest prawdopodobnie zwi¹zane ze s³abszym stopniem zmetamorfizowania wystêpuj¹cych tu tzw. wapieni wojcieszowskich (tzw. epimetamorfizm) w porównaniu do marmurów Masywu nie¿nika, przez co ska³a jest mniej zwarta, a wêglan wapnia ³atwiej dostêpny. 10. W wietle przedstawionych wyników lichenobiota badanych obszarów wapiennych w Sudetach jawi siê jako bardzo interesuj¹ca, bogata i ró¿norodna. Równoczenie jednak biota ta jest wspó³czenie bardzo zagro¿ona, g³ównie z powodu wystêpowania wiêkszoci odnalezionych gatunków na pojedynczych stanowiskach. Czynniki zagro¿enia dzia³aj¹ na ni¹ zarówno w sposób poredni (g³ównie przez zanieczyszczenie powietrza), jak i bezporedni. Wiêkszoæ analizowanych obiektów stanowi¹ bowiem nieczynne kamienio³omy, które w dobie kryzysu gospodarczego i znacznego bezrobocia, po³¹czonych z du¿ym zapotrzebowaniem na materia³ skalny, mog¹ byæ ponownie uruchamiane. W zwi¹zku z tym nale¿a³oby kilka najcenniejszych pod wzglêdem lichenologicznym obiektów otoczyæ ochron¹ jako stanowiska dokumentacyjne. 50 Maria-Kossowska 50 Appendix 1 List of calcicolous lichen species found in the Polish part of the Sudety Mts The list contains all the calcicolous lichen species found to date in both study areas in the Western and Eastern Sudety Mts. The name of each species is followed by synonyms under which the species has also been reported and the list of localities. Designation of the present-day localities is given in chapter 2. For species found during the present survey brief information on habitat preferences is given, together with the total number of localities (in brackets, separately for each of the study areas). Species not rediscovered today and historical localities of the species are printed in small type. The list is supplemented by maps showing the distribution of each lichen species in the study area according to the ATPOL grid square system. Abbreviations: W.S. Western Sudety Mts; E.S. Eastern Sudety Mts; * non-lichenized fungus; Lit. historical data. Acarospora cervina A.Massal. (Fig. 8.a). On exposed, usually insolated rock faces in quarries. W.S.(5): 6, 12, 16, 20, 26; E.S.(1): 45. Acarospora glaucocarpa (Ach.) Körb. [= Zeora cervina (Pers.) Flot. var. glaucocarpa Wahlenb.] (Fig. 8.b). On exposed rock surfaces in places that are somewhat more shaded than for the preceding species. W.S.(2):, 1, 20; E.S.(3): 36, 41, 45. Lit.: Species not rare on limestones, occuring in the Sudety Mts form the foothills to high mountains (Stein 1879); W.S.: Dziwiszów – Kapela rise, Wojcieszów – Mi³ek Mt. and Po³om Mt. (Flotow 1850). Acarospora heppii (Nägeli in Hepp) Nägeli (Fig. 8.c). Lit.: E.S. Environs of Bystrzyca K³odzka (Stein 1879). Acarospora macrospora (Hepp) A.Massal. ex Bagl. [= A. squamulosa Th.Fr.] (Fig. 8.d). On the top part of exposed marble rock, in extremly insolated place. W.S.: – ; E.S. (1): 43. Lit.: W.S.: Mys³ów – Stary Wapiennik rise, Wojcieszów – Po³om Mt. (Eitner 1901). Acrocordia conoidea (Fr.) Körb. (Fig. 8.e). Lit. E.S.: O³drzychowice Sêdzisz Mt. (Eitner 1901). 51 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 51 Fig. 8. Distribution of the calcicolous lichen species in the Sudety Mts 1946 1970; after 1970 till 1945; a. Acarospora cervina; b. Acarospora glaucocarpa; c. Acarospora heppii; d. Acarospora macrospora; e. Acrocordia conoidea; f. Agonimia gelatinosa 52 Maria-Kossowska 52 Fig. 9. Distribution of the calcicolous lichen species in the Sudety Mts 1946 1970; after 1970 till 1945; a. Arthonia lapidicola; b. Arthopyrenia inconspicua; c. Aspicilia calcarea; d. Aspicilia contorta subsp. contorta; e. Aspicilia contorta subsp. hoffmanniana; f. Aspicilia coronata 53 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 53 Agonimia gelatinosa (Ach.) Brand & Diederich [= Polyblastia gelatinosa (Ach.) Th.Fr.] (Fig. 8.f). On calcareous soil at the bottom of a quarry. W.S.(1): 20; E.S.: – . Arthonia lapidicola (Taylor) Branth & Rostr. [= Coniangium fuscum A.Massal., C. rupestre Körb.] (Fig. 9.a). On east-facing vertical wall in a quarry. W.S.(1): 12; E.S.: – . Lit.: Species common in the whole area of Silesia (Eitner 1896); W.S.: Environs of Lwówek l¹ski (Stein 1879); E.S.: Dolina Kamienicy valley (Fabiszewski 1968). Arthopyrenia inconspicua J.Lahm (Fig. 9.b). On calcareous walls and boulders, in shady places. W.S.(2): 5, 10; E.S.: – . Aspicilia calcarea (L.) Mudd [= A. calcarea L. var. concreta Schaer., Lecanora calcarea (L.) Sommerf.] (Fig. 9.c). On calcareous rocks, usually in extremly insolated, south-faced places. W.S.(3): 13, 22, 26; E.S.: – . Lit.: Acc. Körber (1865) this species is frequent throughout the tegion; W.S.: Boles³awiec (Eitner 1901), E.S.: Czerwoniak Mt. near Krosnowice K³odzkie, environs of Bierkowice (Eitner 1901), Dolina Kamienicy valley, Rogó¿ka (Fabiszewski 1968). Aspicilia contorta (Hoffm.) Kremp. – subsp. contorta incl. A. viridescens (A.Massal.) Hue [= A. calcarea L. var. contorta Flörke, Lecanora contorta (Hoffm.) J.Steiner., Lecanora viridescens (A.Massal.) Müll.Arg.] (Fig. 9.d). On calcareous rocks, in places with a wide range of habitat conditions: horizontal and vertical rock surfaces, in shady or insolated places, occasionally also on limy soil (locality no. 20). Pretty often occurs in the sterile form. W.S.(13): 5, 11, 13, 16, 17, 18, 19, 20, 22, 23, 24, 26, 27; E.S.(6): 28, 32, 33, 34, 35, 41. Lit.: Acc. Körber (1865) this species is widespread throughout the region; W.S.: Wojcieszów Mi³ek Mt. (Eitner 1901); E.S.: Czerwoniak Mt. near Krosnowice K³odzkie (Eitner 1901), Rogó¿ka Wapnisko Mt. (Fabiszewski 1968). Note: Some of the specimens represent an intermediate form between A. contorta subsp. contorta i A. contorta subsp. hoffmanniana described by Ekman i Fröberg (1988), characterized by darker thallus. 54 Maria-Kossowska 54 – subsp. hoffmanniana Ekman & Fröberg [= Lecanora hoffmannii (Ach.) Flagey, Aspicilia calcarea L. var. hoffmanni (Ach.) Stein] (Fig. 9.e). On limestones in exposed, usually south-facing places. W.S.(5): 13, 18, 20, 22, 26; E.S.: – . Lit.: W.S.: Mojesz Dolny (Stein 1889), Radomice – Wapienica rise, Siedlêcin Wapienna Mt. (Eitner 1901); E.S.: ¯elazno Wapniarka Mt. (Eitner 1901). Aspicilia coronata (A.Massal.) Anzi [= Lecanora cornuligera Zahlbr., L. coronata (A.Massal.) Jatta] (Fig. 9.f). On exposed rock, in insolated place. W.S.(1): 13; E.S.: – . Lit.: W.S.: Wojcieszów Mi³ek Mt. (Eitner 1901); E.S.: Jaszkowa Górna near K³odzko (Eitner 1901). Aspicilia farinosa (Flörke) Arnold [= A. calcarea L. for. farinosa Flörke, Lecanora farinosa (Flörke) Arnold] (Fig. 10.a). On rock faces with a western exposure. Only sterile specimens found. W.S.(1): 23; E.S.: – . Lit.: W.S.: Wojcieszów Mi³ek Mt. (Eitner 1901). Aspicilia moenium (Vain.) G.Thor & Timdal (Fig. 10.b). On calcareous walls and boulders in exposed places, occasionally also on limy soil (locality no. 20). Usually forms only single squamules. W.S.(4): 1, 12, 20, 26; E.S.: – . Bacidia bagliettoana (A.Massal. & De Not.) Jatta [= Bacidia muscorum (Sw.) Mudd] (Fig. 10.c). On mosses, plant debris and directly on the limy soil, in insolated places. W.S.(3): 12, 22, 26; E.S.(1): 43; Bacidia trachona (Ach.) Lettau [= Bilimbia coprodes Körb.] (Fig. 10.d). Lit.: E.S.: Rogó¿ka (Eitner 1901). Bacidina egenula (Nyl.) Vìzda (Fig. 10.e). Lit.: E.S.: ¯elazno – Wapniarka Mt. (Eitner 1896). Bilimbia lobulata (Sommerf.) Hafellner & Coppins [= Mycobilimbia lobulata (Sommerf.) Hafellner, Toninia lobulata (Sommerf.) Lynge, T. syncomista Flörke] (Fig. 10.f). On calcareous soil covered by mosses, in sun-exposed place. W.S.: – ; E.S.(1): 32. Lit.: E.S.: O³drzychowice Sêdzisz Mt. (Eitner 1901); Rogó¿ka (Fabiszewski 1968). 55 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 55 Fig. 10. Distribution of the calcicolous lichen species in the Sudety Mts 1946 1970; after 1970 till 1945; a. Aspicilia farinosa; b. Aspicilia moenium; c. Bacidia bagliettoana; d. Bacidia trachona; e. Bacidina egenula; f. Bilimbia lobulata 56 Maria-Kossowska 56 Bilimbia sabuletorum (Schreb.) Arnold (Fig. 11.a). On saxicolous and terricolous mosses, in one case also directly on rock surface (locality no. 45), in insolated or shady places. W.S.(5): 1, 9, 19, 22, 26; E.S.(2): 41, 45. Caloplaca alociza (A.Massal.) Migula [= Callopisma agardhianum Ach.] (Fig. 11.b). Lit.: E.S.: O³drzychowice Sêdzisz Mt., ¯elazno Wapniarka Mt. (Eitner 1901). Caloplaca cerina (Ehrh. ex Hedw.) Th.Fr. – var. cerina (Fig. 11.c) On dolomitic rock face with a south-eastern exposure, in insolated place. W.S.: – ; E.S.(1): 32. Lit.: W.S.: Wojcieszów – Po³om Mt. (Stein 1879). – var. muscorum A.Massal. [= C. stillicidiorum (Vahl) Lynge, Callopisma cerinum (Ehrh.) De Not. for. stillicidiorum Ach., Callopisma cerinum var. chloroleuca (Sm.) Stein, Zeora cerina var. chloroleuca E.Bot.] (Fig. 11.d). On mosses and plant debris in close proximity to limestone blocks, in insolated place. W.S.(1): 22; E.S.: – . Lit: Stein (1879) mentioned C. cerina var. muscorum as species occuring here and there in the mountains; acc. Eitner (1901) it was widespread in the region; W.S.: Wojcieszów Po³om Mt. (Flotow 1850), Mojesz (Stein 1879). Caloplaca chalybaea (Fr.) Müll.Arg. (Fig. 11.e). On limestone rocks and boulders. W.S.(3): 5, 13, 20; E.S.: – . Lit.: W.S.: Mys³ów Stary Wapiennik rise, Wojcieszów – Po³om Mt. and Mi³ek Mt. (Eitner 1901); E.S.: Stronie l¹skie Krzy¿nik Mt. (Eitner 1901). Caloplaca cirrochroa (Ach.) Th.Fr. [= Amphiloma cirochroum (Ach.) Körb., A. murorum Hoffm. var. cirrochroum Ach., Gasparinia cirrochroa Ach., Placodium murorum Hoffm. var. cirrochroum (Ach.) Sch.] (Fig. 11.f). On limestones, in shady and insolated places. W.S.(4): 2, 10, 13, 18; E.S.: – . Lit.: W.S.: Wojcieszów – Mi³ek Mt. and Po³om Mt. (Flotow 1850, Körber 1855, 1865, Eitner 1911), Mys³ów Stary Wapiennik rise (Eitner 1911). 57 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 57 Fig. 11. Distribution of the calcicolous lichen species in the Sudety Mts 1946 1970; after 1970 till 1945; a. Bilimbia sabuletorum; b. Caloplaca alociza; c. Caloplaca cerina var. cerina; d. Caloplaca cerina var. muscorum; e. Caloplaca chalybaea; f. Caloplaca cirrochroa 58 Maria-Kossowska 58 Caloplaca citrina (Hoffm.) Th.Fr. (Fig. 12.a). On calcareous rocks, in places with a wide range of habitat conditions: horizontal and vertical rock surfaces, in deep shade as wellaes in fully insolated places. W.S.(10): 1, 2, 5, 8, 12, 13, 17, 21, 22, 23; E.S.(5): 28, 29, 34, 40, 41. Lit.: Not rare in the whole area of Silesia from the lowland to mountains (Stein 1879). Caloplaca coronata (Kremp. ex Körb.) J.Steiner (Fig. 12.b). On limestone rock in fully insolated place. W.S.(1): 13; E.S.: – . Caloplaca crenulatella (Nyl.) H.Olivier (Fig. 12.c). On rock faces in quarries, in insolated places. W.S.(3): 12, 20, 22; E.S.: – . Caloplaca decipiens (Arnold) Blomb. & Forsse (Fig. 12.d). On clacareous faces and boulders, in insolated places. W.S.(4): 13, 20, 22, 26, E.S.: – . Lit.: E.S.: Rogó¿ka Wapnisko Mt. (Fabiszewski 1968). Caloplaca erythrocarpa (Pers.) Zwackh (Fig. 12.e). Lit.: E.S.: K³odzko County (Stein 1879). Caloplaca holocarpa (Hoffm. ex Ach.) A.E.Wade [= Caloplaca lithophila H.Magn.] (Fig. 12.f). On limestone walls, usually in exposed places. W.S.: – ; E.S.(3): 33, 35, 42. Lit.: Acc. Stein (1879) species common on various habitats in the whole area of Silesia. Caloplaca lactea (A.Massal.) Zahlbr. (Fig. 13.a). On vertical rock surfaces, in exposed places. W.S.(1): 12; E.S.(2): 28, 41. Lit.: E.S.: Dolina Kamienicy valley (Fabiszewski 1968) Caloplaca neglecta (Körb.) Migula [= Catillaria neglecta Körb.] (Fig. 13.b). Lit.: W.S.: Wojcieszów – Po³om Mt. (Eitner 1901) 59 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 59 Fig. 12. Distribution of the calcicolous lichen species in the Sudety Mts 1946 1970; after 1970 till 1945; a. Caloplaca citrina; b. Caloplaca coronata; c. Caloplaca crenulatella; d. Caloplaca decipiens; e. Caloplaca erythrocarpa; f. Caloplaca holocarpa 60 Maria-Kossowska 60 Caloplaca saxicola (Hoffm.) Nordin [= Caloplaca murorum (Hoffm.) Th.Fr.] (Fig. 13.c). On vertical, south-facing wall in a quarry, in place sheltered by young trees. W.S.: – ; E.S.(1): 29. Lit.: Species common throughout Silesia from the lowland to the mountains, on walls, rocks and boulders (Stein 1879); E.S.: Dolina Kamienicy valley (Fabiszewski 1968). Caloplaca teicholyta (Ach.) J.Steiner (Fig. 13.d). On calcareous rocks in exposed places. W.S.(1): 22; E.S.(1): 33. Caloplaca variabilis (Pers.) Müll.Arg. [= Zeora variabilis (Pers.) Flot., Callopisma variabile Pers.] (Fig. 13.e). On calcareous rocks, in insolated and shady places. W.S.(3):10, 13, 24; E.S.: – . Lit.: Acc. Stein (1879) and Eitner (1901) this species was widespread in the hills and mountains of Silesia; W.S.: Dziwiszów – Kapela rise, Nielestno near Wleñ, Wojcieszów Mi³ek Mt. (Flotow 1850); E.S.: K³odzko County (Flotow 1850), ¯elazno Wapniarka Mt. (Eitner 1911). Caloplaca velana (A.Massal.) Du Rietz (Fig. 13.f). On rock faces and blocks, usually in exposed, insolated places. W.S.(5): 7, 20, 22, 26, 27; E.S.(3): 41, 42, 43. Candelariella aurella (Hoffm.) Zahlbr. [= Zeora vitellina Ehrh. var. aurella (Hoffm.) Flot.] (Fig. 14.a). On rock faces and boulders, in more or less exposeed places. W.S.(12): 1, 7, 8, 11, 12, 13, 17, 18, 20, 22, 26, 27; E.S.(7): 32, 33, 34, 38, 41, 42, 43. Lit.: W.S.: Dziwiszów – Kapela rise, Wojcieszów – Po³om Mt. (Flotow 1850); E.S.: Kletno (Fabiszewski 1968). Candelariella vitellina (Hoffm.) Müll.Arg. (Fig. 14.b). On calcareous rocks (!), in exposed places. W.S.(2): 13, 22; E.S.: – Note: Species typical for the siliceous rock substrates. 61 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 61 Fig. 13. Distribution of the calcicolous lichen species in the Sudety Mts 1946 1970; after 1970 till 1945; a. Caloplaca lactea; b. Caloplaca neglecta; c. Caloplaca saxicola; d. Caloplaca teicholyta; e. Caloplaca variabilis; f. Caloplaca velana 62 Maria-Kossowska 62 Catapyrenium cinereum (Pers.) Körb. (Fig. 14.c). Lit.: W.S.: Wojcieszów – Po³om Mt. (Körber 1855). Catapyrenium daedaleum (Kremp.) Stein (Fig. 14.d). Lit.: W.S.: Wojcieszów – Po³om Mt., Mojesz Górny (Stein 1879). Catillaria chalybeia (Borrer) A.Massal. (Fig. 14.e). On limestones in insolated places. W.S.(4): 5, 7, 12, 26; E.S. (2): 32, 43. Catillaria lenticularis (Ach.) Th.Fr. [= Biatorina lenticularis Ach., Blastenia lenticularis Flot.] (Fig. 14.f). On calcareous rocks, in fully insolated as well as more of less shaded places. W.S. (7): 13, 20, 21, 22, 24, 26, 27; E.S. (5): 28, 29, 32, 33, 43. Lit.: W.S.: Wojcieszów – Po³om Mt. and Mi³ek Mt. (Flotow 1850), Siedlêcin Wapienna Mt. (Eitner 1901). Cladonia glauca Flörke (Fig. 15.a). On mosses and limy soil. W.S. (1): 22; E.S.: – . Cladonia pocillum (Ach.) Grognot [= Cladonia pyxidata (L.) Fr. var. pocillum (Ach.) Flot.] (Fig. 15.b). On limy soil and rock surfaces covered by mosses. W.S.(5): 7, 19, 22, 24, 26; E.S.(3): 33, 34, 45. Lit.: Stein (1879) reported all forms of C. pyxidata (including f. pocillum) as common in the Sudety Mts. Clauzadea monticola (Schaer.) Hafellner & Bellem. [= Lecidea monticola (Ach.) Ach., L. fuscorubens Nyl., Lecidella ochracea Hepp] (Fig. 15.c). On limestones in places with different degrees of an insolation. W.S.(3): 5, 13, 23; E.S.(4): 36, 39, 43, 45; Lit.: W.S.: Wojcieszów – Mi³ek Mt. (Körber 1865), Wojcieszów – Po³om Mt. (Eitner 1901); E.S.: Rogó¿ka, Stronie l¹skie Krzy¿nik Mt. (Eitner 1901). 63 Lichens-growing-on-calcareous-rocks-in-the-Polish-part-of-the-Sudety-Mountains 63 Fig. 14. Distribution of the calcicolous lichen species in the Sudety Mts 1946 1970; after 1970 till 1945; a. Candelariella aurella; b. Candelariella vitellina; c. Catapyrenium cinereum; d. Catapyrenium daedaleum; e. Catillaria chalybeia; f. Catillaria lenticularis