Deposits and fauna of the Sudetic caves – the state of research
Transcription
Deposits and fauna of the Sudetic caves – the state of research
Deposits and fauna of the Sudetic caves – the state of research Jerzy Bieroński1, Paweł Socha2, Krzysztof Stefaniak3 1 Department of Physical Geography, Institute of Geography and Regional Development, University of Wrocław, Pl. Uniwersytecki 1, 50-137 Wrocław, e-mail: jbier@osti.pl 2 Department of Paleozoology, Zoological Institite, University of Wrocław, ul. Sienkiewicza 21, 50-335 Wrocław, e-mail: socha@biol.uni.wroc.pl 3 Department of Paleozoology, Zoological Institite, University of Wrocław, ul. Sienkiewicza 21, 50-335 Wrocław, e-mail: stefanik@biol.uni.wroc.pl ABSTRACT Karstic areas in the Sudetes occur mainly in two regions: Kaczawskie Mts and Eastern Sudetes. In this areas mostly metamorphic carbonate rocks undergo karstification. The Sudetic caves formed since Palaeogene till recent times. However, documented cave deposits are referred to the Neogene till Holocene. The paper contains a review of German and Polish research on the deposits and animal remains from the Sudetic caves. A considerable part of the deposits was redeposited which renders their interpretation difficult. Dating of dripstones indicates that they were formed at the end of the middle and in the upper Pleistocene. It was found that most bone remains come from the Vistula glaciation which was confirmed by few datings. Only in the case of Jaskinia Południowa (Mt. Połom, Kaczawskie Mts) there is information on a single finding of a Pliocene rodent Baranomys langenhani - probably B. loczyi. The few earlier mentions of the presence of Palaeolithic man in the Sudetic caves in most cases (except caves of Mt. Połom) were not confirmed. KEY WORDS: palaeography, palaeoecology, caves, caves deposits, Sudetes Mts., Poland Introduction Karstic caves of the Sudetes are associated with outcrops of carbonate rocks forming intercalations of various size among non-karstifying rocks. The intercalations or larger bodies of carbonate rocks are grouped in several regions in the Sudetes (Fig. 1). They occupy the largest areas in the Western Sudetes – in the Kaczawskie Mts. (IA) and in the neighbouring Pogórze Bolkowskie (IB). The second karstic area of the Sudetes is the margin of the Kotlina Kłodzka – the massif of Mt. Śnieżnik (II), the range of Krowiarki (III), Złote Mts. (IV) and Bystrzyckie Mts. (V). Smaller outcrops of calcareous rocks are found also in other massifs. The German studies in the Kaczawskie Mts focused on the region of Wojcieszów. The caves explored in Mt. Połom included Wschodnia Cave (Eastern Cave, Hellmich- höhle), Północna Duża Cave (Big Northern Cave, Witschelhöhle) and Południowa Cave (Southern Cave, Kitzelhöhle, Kitzelloch, Kitzelberghöhle) (Fig. 1; IA), the rock shelters on Mt. Miłek – Cisowe I Rock Shelter and Cisowe II Rock Shelter (Yew Tree Shelter I and II, Eibenloch I, Eibenloch II) (Fig. 1; IB). In the Eastern Sudetes the studies included Rogóżka Cave (Cave in Rogóżka, Wolmsdorferhöhle, Wolmsdorfer Tropfsteinhöhle) (Fig. 1; III) and Radochowska Cave (Reyersdorfer Tropfsteinhöhle, Reyersdorfer Höhle) (Fig. 1; IV). The exploration yielded fossil faunistic material and traces of human occupation. The studies before World War II are at present difficult to verify precisely (damage of caves or lost fossil record). 184 Jerzy Bieroński, Paweł Socha, Krzysztof Stefaniak Fig. 1. Location of karstic areas in the Polish Sudetes (after Pulina 1989, modified). IA – Kaczawskie Mts.; IB – Pogórze Bolkowskie; II – Massiif of Mt. Śnieżnik; III – Krowiarki Range; IV – Złote Mts.; V – Bystrzyckie Mts. Kaczawskie Mountains Mount Połom A profile of deposits composed of several layers of loams ranging in colour from red to brown, with intercalations of quartz gravel and fragments of dripstones and limestone was described in Wschodnia Cave. Heller (1937) and Zotz (1939) described two faunistic assemblages. The age of the older assemblage was estimated as early Pleistocene (Cromerian), that of the younger assemblage as upper Pleistocene (probably starting with the Eem interglacial). The reinterpretation of the results of German workers done by Kowalski (1954), indicates an Upper Pleistocene and Holocene age of the bone remains (the material includes no species which would be characteristic for older periods of Pleistocene). Our exploration of the localities on Mt. Miłek revealed a similar faunal assemblage in the deposits of rock shelters – among others Małgorzata Rock Shelter and Trwoga Paleontologa Rock Shelter (Palaeontologist’s Fright Rock Shelter); its age was estimated as Upper Paleistocene to Atlantic period of the Holocene (Pakiet 1999). The species composition (Table 1) of the assemblage indicates a forest environment (beechwood-type deciduous forests and mixed forests) and proximity of water courses. Two asemblages of bone breccia were found before World War II in Południowa Cave in two different locations (near the entrance and inside the cave). The discovery of the new species of hamster Baranomys langenhani in that cave (bone breccia near the cave entrance) is noteworthy. At presence the species is referred to as B. loczyi KORMOS, 1933 (Nadachowski 1989). The species represents the genus Baranomys KORMOS, 1933 which became extinct in the Villanian. The opinions of the German researchers on the age of these bone remains differed. Zotz (1937a, 1939) postulated that they were of the same age, while Heller (1937) indicated a different age. It can be supposed that the breccia from Południowa Cave and the terra rossa-type deposits which are present in most caves of the area were formed in the Miocene and Pliocene in conditions of a warm, humid climate in a mostly woodland environment. However, the presence of the genus Baranomys indicates the presence of cooler, arid, open areas (forest-steppe, savanna). 185 Deposits and fauna of the Sudetic caves – the state of research Table 1. Mammal remains found in caves of Mt. Połom before World War II. Takson Wschodnia Cave Północna Cave Południowa Cave Talpa europea L. Soricidae indet. Myotis bechsteini (K.) Myotis dasycneme (B.) Rhinolophus aff. ferrum equinum Myotis sp. Lepus sp. Baranomys langenhani H. Sciurus vulgaris L. Arvicola terrestris (L.) Glis glis (L.) Ursus arctos (L.) Ursus spelaeus R. et H. Crocuta spelaea (G.) Felis sp. Panthera spelaea (G.) Martes sp. Martes cf. foina (E.) Mustela putorius L. Equus sp. Rangifer tarandus (L.) Ungulata indet. + + + + + + + + + + - + + + + + + + + + + - Caves and rock shelters of the CracowWieluń Jura commonly contain deposits and breccias of a similar type. It can be supposed that the presence of diverse and much higher situated karst created conditions for the development of a mosaic habitat. Higher situated areas of limestone rocky substratum provided good conditions for the development of vegetation and fauna, characteristic of dry and warm open areas or park-type forests. Most probably lower situated areas in waterrich deep valleys were covered in forests of diverse character (Głazek & Szynkiewicz 1985, Kowalski 1989, 1990, Stefaniak 1995, 2004, Croitor & Stefaniak in press). Animal bones and artefacts were found in Północna Duża Cave (Table 1) (Zotz 1937, 1939). The deposits of the cave are Pleistocene cave loams (with Upper Pleistocene mammalian bones), layers of calcite dripstone covers and breccia similar to those found in Południowa Cave (Kowalski 1954). Near Jaskinia Wschodnia another cave was situated, called Kammerberghöhle. It was Obok Wschodniej Cave + + + + + + + + + - destroyed during quarrying and its exact location is still uncertain. On the dump near this cave In 1926 Wenke found flint tools, and Zotz and quarry workers – very numerous mammal bones (Zotz 1939, Kowalski 1954), also some bearing traces of human processing. Bones of the cave bear and other Pleistocene and Holocene mammals were the most numerous. The faunal remains, like in other caves of the area, were deposited in various periods of the Vistula glaciation and Holocene. Based on available data, it can be said that the bone remains came from the last stages of the Vistula glaciation and the beginning of Holocene. This is indicated by fluoro-apatite datings of the bones (Pulina 1977) from Naciekowa Cave: 25-45 kA - Grudziądz, Hengelo, Denekamp interstadials, main stadial of the Vistula glaciation. Two main periods of deposition of bone remains and sediments were distinguished in the caves of Mt. Połom. The oldest phase, starting in the Palaeogene and continuing till the Pliocene, is represented by calcite breccias Table 2. Mammal remains found in deposits of rock shelters of Mt. Miłek. TAKSON Erinaceus sp. Sorex araneus L. Chiroptera indet. Eptesicus cf. serotinus (S.) Lepus sp. Lepus europaeus (P.) Lepus timidus P. Oryctolagus cuniculus (L.) Ochotona sp. Ochotona pusilla P. Sciurus vulgaris L. Dicrostonyx gulielmi (S.) Lemmus lemmus (L.) Clethrionomys glareolus (S.) Arvicola sp. Arvicola terrestris (L.) Pitymys sp. Microtus sp. Microtus agrestis (L.) Microtus arvalis (P.) Microtus subterraneus (d. S-L.) Microtus oeconomus (P.) Microtus (Stenocranius) gregalis (P.) Cricetidae indet. Cricetus cricetus (L.) Apodemus sylvaticus (L.) Glis glis L. Canis lupus L. Ursus sp. Ursus arctos (L.) Vulpes vulpes (L.) Mustela erminea L. Martes sp. Małgorzata Rock Shelter + + + + + + + + + + + + + + + + + + + - Lucia Rock Shelter + + + + + + + + + - Trwoga Paleontologa Rock Shelter + + + + + + + + + + + + + + + + - Tomkowa Niche + + + + + + + + + + - Panna Rock Sheltr + + + + + + + + + - Cisowe 1 Rock Shelter + + + + + + + + + + + + + + + + + Cisowe 2 Rock Shelter + + + + + + + + + Table 2. Mammal remains found in deposits of rock shelters of Mt. Miłek – contitnation from preceding page. TAKSON Mustela nivalis L. Mustela putorius L. Felis silvestris S. Meles meles (L.) Martes martes (L.) Equus caballus (L.) Sus scrofa L. Cervus elaphus L. Capreolus capreolus (L.) Małgorzata Rock Shelter + + + + + + Lucia Rock Shelter + - Trwoga Paleontologa Rock Shelter + + + + + + Tomkowa Niche + - Panna Rock Sheltr + - Cisowe 1 Rock Shelter + + + + + Cisowe 2 Rock Shelter + + + + - dripstones, breccias of crystalline rocks from Południowa Cave and Północna Cave, as well as red loams of „terra rossa” type, present in most caves of this area. During the Pleistocene cave loams were deposited in these caves, and sometimes also gravels. Bone remains were present in Tertiary breccias and Pleistocene and Holocene cave loams. Neogene red loams also contain bone remains which got there as a result of redeposition during the Pleistocene and Holocene. In the Pleistocene and on the Pleistocene/Holocene boundary in the caves of Mt. Połom sediments with bone remains were redeposited. The reasons could be both gravitational transport processes and flowing or infiltrating waters. For this reason conclusions of the German authors regarding the age of the fauna, older than the Vistula glaciation, are doubtful. 3 m thick (Pakiet 1999). Among gastropods, 49 taxa were identified. Palaeoecological analysis showed the presence of shade-loving forest forms in the whole profile; open-country species were numerous in the bottom part of the profile; mesophile forms increased in dominance with decreasing depth. The age of the vertebrate and snail remains was estimated as the period from late Pleistocene till Holocene (Pakiet 1999). Lucia Rock Shelter is funnel-like, narrowing downwards. The profile is ca. 0.5 m wide and ca. 1 m deep. Based on the malacological analysis the deposits from Lucia Rock Shelter (35 snail species) were dated as middle and upper Holocene (Pakiet 1999). Mount Miłek Studies on rock shelters on Mt. Miłek (Fig. 1; IB) were initiated by Zotz (1939). He studied two such shelters: Cisowe I (length 5 m) and II (length 3 m). The material obtained from the shelters included remains of snails and vertebrates (Tab. 1). Besides, charcoal and remains of a campfire were found. The age of the deposits in the studied rock shelters was estimated as Vistulian glaciation – Cisowe I and Holocene – Cisowe II (Zotz 1939). Our post-war studies in the Kaczawskie Mts focused on Mt. Miłek. Deposits of the following rock shelters were explored: Małgorzata Rock Shelter (Małgorzata’s Rock Shelter), Lucia Rock Shelter (Lucia’s Rock Shelter), Trwoga Paleontologa Rock Shelter, Tomkowa Niche (Tomek’s Niche), Panna Rock Shelter (Maiden Rock Shelter). The rock shelters are grouped around the outcrop Cisowe, near its top and below. Numerous remains of snails and vertebrates (Table 2) were found in all the sites. The most numerous remains were found in Małgorzata Rock Shelter (Fig. 2). The shelter is located near Rock Shelter Cisowe I and includes a rock niche and a profile of deposits situated below, Fig. 2. Site profile and sediments of Małgorzata Rock Shelter (after M. Pakiet 1999, modified). 1. marble fragments; 2. soil; 3. silts; 4. silts with limestone lumps; 5. sandy loams with limestone lumps; 6. sandy loams; 7. silty loams; 8. sand; 9. marbles; 10. sampling points for malacospectra. Deposits and fauna of the Sudetic caves – the state of research Trwoga Paleontologa Rock Shelter is located on a rock face and has the form of a niche (ca. 1 m long). Its deposits consist of dusty loam with fragments of limestone and sandy loams. The character of the deposits indicates their sliding from higher-situated crevices. Fourteen species of snails were found in the deposits, with the dominance of taxa characteristic for partly shaded and mesic habitats. The time of deposition of the remains can be estimated as the Atlantic period and late 189 Holocene (Pakiet 1999). Loamy and dusty deposits (depth 20 cm) of Tomkowa Niche contained forest snails of shaded and mesic habitats, representing forms of middle and late Holocene (Pakiet 1999). The last explored site was Panna Rock Shelter. The shelter is a small rock niche at the base of Cisowe rock. Malacofaunistic studies of Pakiet (1999) indicate that the deposits were formed during the Holocene. Eastern Sudetes Krowiarki Range German studies in the Krowiarki Range included Rogóżka Cave (Wolmsdorferhöhle) (Fig. 1; III). Rogóżka Cave was located near Konradów in the region of Lądek Zdrój, in a marble quarry ”Rogóżka”, with the entrance on the lowest mining level, about 20 m above the bottom of the Konradka stream valley. In their studies Pax and Zotz found that the deposits were composed of yellow-brown cave loams (Pax & Maschke 1935, Pax 1937, Zotz 1939). They were practically devoid of faunal remains and human artefacts, except for two bird bones and a fragment of spruce timber (Pax 1937, Pax & Maschke 1935). The earlier reported finding of fossil fauna in the deposits of that cave was regarded as doubtful already Fig. 3. Map of the Na Ścianie Cave (after M. Pulina 1996). Plan in left corner shows part of the cave situated below Sala Złomisk. by the German authors. An interesting and rich extant fauna was also found in that cave and studied by Arndt (1921, 1923), Seidel (1927), Pax & Maschke (1935), Pax (1936b, 1937) and others. More than 70 animal species were found there, including one new species and two new varieties (Kowalski 1954, Pulina 1996). Another cave situated in the same quarry is Na Ścianie Cave (On-The-Wall Cave) (Fig. 3). Its entrance is located on the headwall of the quarry, 41 m above the bottom of the lowest mining level (c. 670 m a.s.l.). The length of the corridors is about 250 m, and the total denivelation is 21 m. The cave was discovered in 1985 by explorers from Stronie Śląskie (Pulina 1996) and studied immediately after its discovery (Bieroński & Wiszniowska 1994). The subterranean fauna was studied by Pomorski (1987). Na Ścianie Cave is unique among the caves of the region due to the great variety of its deposits (including dripstones) and its high location above the valley bottom. Among identified deposits of that cave the following can be distinguished: two dripstone formations of different age (older and younger, the latter probably Holocene), clayey siphonal deposits, sandy deposits of flowing water and block screes resulting from subsidence of the roof rocks. Further sequences of deposits associated 190 Jerzy Bieroński, Paweł Socha, Krzysztof Stefaniak with the cave are found near the entrance, on the quarry wall: silty filling of the fossil karst doline and a complex of varied deposits filling a fossil cave corridor (Fig. 4). The latter contains a fairly thick dripstone cover formed on the wall of the original corridor, mostly cracked into fragments as a result of mining (Fig. 4; roof of the profile). Later the corridor became completely filled with a sandy deposit with admixture of gravel brought by flowing water. The deposit was then cemented with calcite, to form cave sandstone. Few silty balls and gravel grains were contained in it. Subsequently the sandstone became partly eroded by water, resulting in interesting corrosion-erosion forms. The final stage of formation of this deposit complex was filling the corridor formed in the sandstone with loose sand brought by water and loam and clays of fine lamination. Clayey deposits with silt intercalations were found in the terminal part of the cave – Korytarz Nadziei (Corridor of Hope) (Fig. 3). The strata were disturbed, indicating periodical erosion and subsidence. When the corridor was discovered it had a fossil character. Its terminal part was mostly filled by sandy deposits with clay. Subfossil bat remains: Myotis mystacinus (KUHL, 1819), Myotis nattereri (KUHL, 1818) and an unidentified Myotis sp. were found in the deposits of this part. They indicate the Holocene age of the clayey deposits in the corridor. Considering the incomplete documentation of the deposits of Rogóżka Cave, it is difficult at present to conclude about its origin and development. The cave was located low above the valley bottom of a small tributary of the Konradka stream and could develop as a separate system from the higher situated Jaskinia Na Ścianie. After World War II local people observed formation of karst dolines in the bottom of the valley of a small stream near the quarry. This indicates contemporary active karstic processes. However, formation of the dolines associated with reactivation of older fossil karst cannot be excluded. Such karst could be related to Rogóżka Cave. Na Ścianie Cave undoubtedly documents the stage of development of the Konradka valley bottom in the period when it was located c. 60 m higher than it is now. The existence of fossil corridor with cave sandstone indicates a multi-stage development of the cave and its deposits. However, only the youngest deposits (Holocene) in the siphonal Nadzieja Corridor (Corridor of Hope) have been documented in detail to date. Fig. 4. Profile of the deposits filling fossil part of Na Ścianie Cave (depth of profile in metres – left axis; layer’s number – right axis) (after J. Bieroński, T. Wiszniowska 1994, modified). Explanations: 1. surface of the quarry scarp; 2. blocks of marbles and flowstone; 3. gray-yellowish loam with fragments of marbles, flowstone and flowstone breccias; 4. horizontally oriented marble blocks up to 60 cm diameter; 5. brownyellowish loam with fragments of cave sandstone and cemented marble breccias; 6. blocks of cave sandstone in sandy loam; 7. fine-grained clayey sand; 8. plate of cave sandstone; 9. fine-grained loamy sand; 10. horizontally oriented cave sandstone blocks up to 60 cm diameter; 11. fine, laminated clayey sands; 12. poorly sorted gravels with fragments of rolled clay sediments; 13. fine-grained clayey sands; 14. sands with angular gravels and clay bricks; 15. fine-laminated sandy mud; 16. fine-laminated clay; 17. cave sandstone (thickness not known). Deposits and fauna of the Sudetic caves – the state of research Bystrzyckie Mountains Small outcrops of crystalline limestone are present in the Bystrzyckie Mts. Solna Jama Cave (Salzlöcher, Salt Pit) (Fig. 1; V) is located in one of them, in an abandoned quarry near Gniewoszów (region of Międzylesie). It is situated near two small streams. Its length is 40 m. It was known already in the 18th c. which is indicated by the graffiti on its walls. Till World War II it was known mostly for its interesting extant fauna. After the war mainly faunistic studies were conducted there. A few dozen recent animal species were recorded from the cave (Arndt 1921, Stammer 1936, Stach 1947, Wołoszyn 1968, Skalski 1970, 1976, 1994, Pomorski 1990, 1992). Palaeontological studies in the cave were carried out in 1984-1985 by our team, organised by Teresa Wiszniowska and Marek Pakiet. A niche was found in the cave roof during the studies. The niche ended with a narrow crevice filled by loose breccia with cave bear bones. 191 suggested time of the deposition of bone remains is the end of Pleistocene and beginning of Holocene. The origin of Solna Jama is probably associated with corrosion of crevices by one of the neighboring streams. The lowest situated part of the cave is permanently flooded. During the 1985 studies the largest chamber of the cave was discovered, its floor flooded. The cave is drained by small karst springs located on the valley slope. Discovery of the crevice with remains of the cave bear indicates that the cave must have a higher situated horizon, as yet undiscovered. Złote Mountains Table 3. List of mammal species from deposits of Solna Jama Cave. Takson Sorex minutus L. Crocidura sp. Myotis myotis (B.) Plecotus auritus (L.) Sciurus vulgaris L. Clethrionomys glareolus (S.) Cricetuss sp. Microtus sp. Arvicola terrestris (L.) Ursus spelaeus R. et H. Gulo gulo (L.) Carnivora sp. Most deposits were cave loams. As a result of exploration of the deposits mammal bone remains were found (Table 3). The cave is noteworthy because of the finding of remains of the wolverine Gulo gulo (LINNAEUS, 1758). It is one of the few localities of the species in Poland (Wolsan 1989). The species is characteristic for boreal forests of taiga type. Palaeoecological analysis of the mammal assemblage indicates existence of a taiga-type forest with some streams and meadows in the vicinity of the cave. The Fig. 5. Hipothetic section of galleries in the western part of Radochowska Cave (diagrammatic) (after J. Bieroński et al. 1985, modified). Explanations: 1. dolomitic marbles; 2. slope loams (Pleistocene and Holocene); 3. collapsed roof of the upper level of the cave (?Holocene); 4. residual clay from the upper level (Pleistocene, ?Holocene); 5. local connection between upper and lower level; 6. clays from upper level fulfilled lower level (Pleistocene, Holocene); 7. layer of clays infilated with water; 8. clay with blocks of limestone (Pleistocene); 9. Sala Stołowa; 10. karst doline fordem as a result of substratum settling, situated on the slope; 11. extent of excavation in 1983; 12. extent of excavation of L. Zotz in years of 1935-1936. One of the longest and best known Sudetic caves is Radochowska Cave (Reyersdorfer Tropsteinhöhle) (Fig. 1; IV). It has been known since the 18th c., and three artificial entrances lead to it. It is among the best studied caves of the area (Frenzel 1936, 1937a, 192 Jerzy Bieroński, Paweł Socha, Krzysztof Stefaniak b, Pax 1936 a, b, 1937, Stach 1936, 1947, such a situation the bones present on the rock Stammer 1936, Zotz 1937 a, b, 1939, ledge (including those cemented by calcite) Kowalski 1954, Walczak 1956, 1958, can be regarded as a trace of the former Wołoszyn 1968, Kos 1978, Bieroński et al. position of the roof of the deposits which fill a 1985, Pulina 1996, Buczyński, Rzońca 2007). part of Sala Stołowa. Discovery of deposits Systematic studies by Zotz (1939) provided containing bone remains in the roof of the first information on the deposits and their crevice north of Sala Stołowa indicates an origin, bone remains and artefacts. The results existence of a higher situated horizon of of Zotz’s (1939) studies in Radochowska Cave chambers, not considered earlier. and the above-mentioned caves in the region Table 4. Radochowska Cave. List of mammal remains of Wojcieszów provided him with the basis to obtained in pre-war studies and in the 1980s. formulate the hypothesis of the ”cave bear cult” among Frenzel Zotz Bieroński Takson Palaeolithic people. On a rock 1936 1939 et al. 1985 ledge, Zotz found bones, and in the Talpa europea L. + Rhinolophus hoposideros (B.) ?+ + rock niche, a cave bear skull Myotis myotis (B.) + covered by a rock plate which he Myotis sp. + interpreted as a ritual burial. Plecotus auritus (L.) + During Frenzel’s (1936) and Zotz’s Lepus europaeus P. + (1939) studies 30 mammal taxa Lepus sp. + were found (Table 4), representing Oryctolagus cuniculus + Sciurus vulgaris L. + + Pleistocene (Vistulian glaciation) Castor fiber L. + and Holocene forms. They Cricetuss sp. + + represent various ecological Microtus arvalis (P.) + groups, from steppe-tundra species Microtus sp. + to euryoecious forms and forestArvicola terrestris (L.) + dwellers. Besides, remains of snail Apodemus sp. + Ursus arctos (L.) + shells, fishes (perch?), amphibians Ursus sp. + + (Rana temporaria LINNAEUS, Ursus spelaeus R. et H. + + + 1758) and an unidentified lizard Canis lupus L. ?+ (Lacerta sp.) were found in the Vulpes vulpes (L.) + + deposits. A similar faunal Felis silvestris S. + ?+ + assemblage was described by Crocuta spelaea (G.) + Meles meles(L.) + + + Bieroński et al. (1985) (Table 4). Martes martes (L.) + In the 1980s the pre-war studies Martes sp. + + were verified by Bieroński et al. Mustela erminea L. + (1985). During their studies they Equus caballus (L. + found deposits with bone remains Equus sp. + + in the roof of the crevice extending Coelodonta antiquitati (B.) ?+ Sus scrofa L. + northwards from Sala Stołowa Cervus elaphus L. + + (Table Hall) and discovered a small Cervus sp. + karst doline above that part of the Megaloceros giganteus B. + cave (Fig. 5). The discovery of the Capreolus capreolus (L.) + + deposits in the crevice threw a new Alces alces (L.) + + light on the interpretation of Zotz’s Bison priscus (B.) + Bison sp. + (1939) studies. The results indicate a possibility of gravitational One of the essential dilemmas redeposition towards Sala Stołowa and concerning Radochowska Cave is the subsidence of the complex of deposits in that information about the presence of Palaeolithic chamber under the effect of water erosion. In man in the cave suggested by Zotz (1939). The Deposits and fauna of the Sudetic caves – the state of research results of studies of Bieroński et al. (1985) do not confirm the hypothesis of the presence of man in the cave. The tools mentioned by Zotz could be shaped by natural processes. The differences in the degree of preservation of the bones mentioned by Zotz might result from redeposition from another location. The layer interpreted by him as an ”ancient floor trodden by animals and humans” may represent deposition by infiltration waters which often change the superficial layer of the deposit in this way. The presence of small charcoal fragments in the deposits of Sala Stołowa also requires an explanation. Numerous charcoal fragments were found by Kos (1978) in a pit near the cave. The possibility of their penetration into the cave during redeposition (probably in Holocene) cannot be excluded. Recent studies disprove also the hypthesis of the ”bear cult” among Palaeolithic people (Leroi-Gourhan 1966). Covering of cave bear skull and vertebrae with ”rock plates” which according to Zotz represented a ”burial” can be explained by natural phenomena of sliding of material along the sloping surface of the deposit (Bieroński et al. 1985). Śnieżnik Massif The 1966 discovery of Niedźwiedzia Cave (Bear Cave) in Kletno, near Stronie Śl. (Fig. 1; II) opened a new, interdisciplinary stage of studies on the Sudetic karst. The reason was the presence of bone-rich deposits in the cave. In subsequent years it turned out to be also the largest known karst system in the Sudetes. The cave system, preserved unaltered by man, made it possible to carry out comprehensive multidisciplinary studies. Clastic deposits of Niedźwiedzia Cave were studied by Pulina (1970, 1989), Szarejko (1976), Bosàk (1989) and Kozłowski (1989). They found prevalence of allochthonous deposits represented by Pleistocene, Holocene and recent spelaeofluvial deposits (silts, clays, gravels, sands). Autochthonous deposits in Niedźwiedzia Cave include mainly rubble 193 deposits, cave clays, block screes and dripstones. A part of these deposits underwent redeposition which in places is still active. In our opinion the significance of redeposition in Niedźwiedzia Cave was clearly underestimated in earlier interpretations. Besides the abovementioned studies on clastic deposits, dripstones of Niedźwiedzia Cave were also studied (Zięba 1978, Głazek 1985, Hercman et al. 1995). Unique studies on the dynamins of recent sedimentation of dripstones in the environment of karstic bowls were done by Fabjańska (1987). Studies on the abundant bone detritus of autochthonous deposits in Jaskinia Niedźwiedzia were carried out by Wiszniowska (1967, 1970, 1976, 1978, 1989), Wiszniowska and Kuryszko (1998), and the results of datings were presented by Wysoczański-Minkowicz (1969). Fig. 6. Situation plan of excavations in middle level of Niedźwiedzia Cave. Location of the profiles in the middle horizon of the cave. Palaeontological studies on the cave deposits yielded material of several hundred thousand remains of vertebrates and snail shells. The studies included chambers of the middle horizon of the cave, where profiles I-V were made (Fig. 6), the lower horizons and the profile below the entrance to Miniaturka Cave (Miniature Cave) (Fig. 7), the profile is located ca. 10 m from the tourist entrance to Niedźwiedzia Cave. The bone material served as the basis for the chronoclimatostratigraphic 194 Jerzy Bieroński, Paweł Socha, Krzysztof Stefaniak analysis of the deposits. The remains were dated with flouro-chloro-apatite and collagen methods (Wysoczański-Minkowicz 1969, Wiszniowska 1989). Based on these datings, four warm periods were distinguished in the studied deposits, separated by cool periods. The age of the remains from various layers was estimated as 13,2 to 15,5 kA. Fig. 7. Profile of sediments below the Miniaturka Cave entrance (after Wiszniowska 1996. Explanations: I – IV – number of layer complex; 1. crystalline limestone; 2. speleothem; 3. zones of calcite deposition; 4. sandy sediments; 5. bones; 6. noncarbonate rocks (gneis, shists); 7. ferruginous-manganese sediments; 8. residua clays; 9. blocks of crystalline limestone. The development of Quaternary fauna of the Massif of Śnieżnik and adjacent areas is known only from the Upper Pleistocene. There are no dated localities with fauna from earlier periods. Palaeoecological analysis and analysis of the composition of faunal remains found in the caves of the region make it possible to reconstruct climatic fluctuations within roughly the last 40 kA (Frenzel 1936; Pax 1936a; Zotz 1939; Wiszniowska 1986; Wiszniowska et al. 1996; Pakiet 1999). The studied material obtained during exploration of deposits of the profiles in various parts of the cave included remains of 29 mammal species (Table 5). The extinct species represented faunal assemblage of mammoth steppe. Montane species were also present (chamoix). The chamoix Rupicapra rupicapra (LINNAEUS, 1758), at present a component of montane faunal community, and found for the first time as fossil in the Sudetes, was a characteristic component of the Pleistocene fauna in Poland (Czyżewska 1989). Cave bear remains dominated. The proportion of the remaining vertebrates and snails in the deposits ranged from a fraction of percent to a few percent. The occurrence of a diverse community of herbivores near the cave (besides the remains of primitive bison, red deer and chamoix) is indirectly suggested by the presence in the deposits of many remains of carnivores which, besides the extant species e.g. wolf, fox, pine marten, included the already mentioned extinct carnivores. In some profiles rodents were present, indicating the presence of water courses (beaver, bank vole). The same is indicated by the large mammals occurring there (red deer, roe deer, primitive bison). The presence of large ungulates indicates the absence of thick snow cover in winter. Niedźwiedzia Cave provided a good shelter for animals to winter (cave bear, badger, bats), or to spend there shorter or longer periods (wolf, cave lion, hyena, other carnivores). The great number of remains of juvenile and very young individuals, especially of the cave bear, indicates the role of the cave as a breeding place. Few remains of large herbivores (e.g. primitive bison, chamoix) may represent leftovers of prey of the carnivores. Some bones, especially of the cave bear, show pathological changes resulting from various injuries and illnesses. Such material constitutes c. 10% collection. It was subject to a separate, detailed analysis including radiographic and histological methods and analysis of element composition (Wiszniowska 1989, Wiszniowska et al. 1996, Wiszniowska, Kuryszko 1998). A detailed analysis of species Deposits and fauna of the Sudetic caves – the state of research composition made it possible to state that most mammal remains represented euryoecious species, with a low proportion of forest- 195 dwellers and forms associated with water bodies. Table 5. List of vertebrate species from deposits of Niedźwiedzia Cave. I – V – profiles; L.H. – lower horizon; P.b.J.M. – profile below Jaskinia Miniaturka Species 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11 12 13 14. 15. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29 Aves indet. Sorex araneus L. Crocidura suaveolens L. Myotis myotis B. - nocek duży Myotis bechsteini (K.) Myotis nattereri (K.) Myotis mystacinus (K.) Myotis daubentoni (K.) Myotis brandti (E.) Plecotus auritus (L.) Eptesicus nilssoni (K. et B.) Castor fiber L. - bóbr Clethrionomys glareolus (S.) Arvicola terrestris (L.) Microtus arvalis (P.) Vulpes vulpes (L.) Canis lupus L. Ursus arctos L. Ursus spelaeus R. Crocuta spelaea (G.) Panthera spelaea (G.) Martes martes (L.) Sus scrofa (L.) Cervus elaphus L. Capreolus capreolus (L.) Rupicapra rupicapra (L.) Bison priscus (B.) The age of the bones ranged from 21800 (±1100) to 28900 (-2200, +3100) years in Sala Lwa (Lion Hall), in Korytarz Człowieka Pierwotnego (Primitive Man Corridor) from >38100 to >40000 years, and in the profile below Miniaturka Cave it was 32100 (±1300) years. Dating of bone remains with C-14 method was done in 2002 at the Institute of Physics, Department of Radioscopes, Radiocarbon Laboratory (Gliwice, Poland). They are the first datings of bones from the cave with this method. Their varied age and the low number of datings make it impossible at present to identify the stratigraphic levels which, in addition, occur in different profiles. Most of the bone remains came from the I + + + + + + + + + + + + + + + + + + - II + + + + + + + + + + + + + + + + + + - III + + + + + + + + + + + + + - Profile IV V + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + L.H. + + + + + + + + + + + - P.b.J.M. + + + + + + + + - period of Grudziądz interstadial and the main stadial of the Vistula glaciation. Examination of bone remains and deposits and the results of dating of bones with C-14 method confirm the hypothesis of their redeposition. Redeposition of at least a considerable part of the studied sediments was noticed already at the initial stage of the studies (Pulina 1970), and solifluction transport was indicated as a likely agent. Subsequent studies showed however that also water transport could be involved (Bosák 1989, Wiszniowska et al. 1996; Bieroński 1997). Anyway, the animal remains contained in the deposits are most probably not in situ. Redeposition makes palaeoecological analysis based on bone remains difficult or 196 Jerzy Bieroński, Paweł Socha, Krzysztof Stefaniak even pointless. The species composition indicates habitats associated with cold steppe or steppe-tundra, with a constant presence of water and of forested areas of various character. The profile below Miniaturka Cave (Wiszniowska et al. 1996) deserves a special attention (Fig. 7). It contained numerous remains of snails which made it possible to carry out a detailed analysis of environmental changes. It is the only profile in the system of Niedźwiedzia Cave with a complete sequence of sediments, including also the end of Pleistocene and Holocene. The main part of the profile, including three complexes of strata, documents the end of Pleistocene (Vistula glaciation). These complexes contained mammal bone remains. Shells of 29 snail species were found in the top part of the profile (complex IV, Fig. 7). All the studied samples from that part show a very high proportion of forest-dwelling snails Isognomostoma isognomostoma (SCHRÖT), Oxychilus depressus (STERKI), shade-loving and higrophilous forms: Ena montana (DRAP.), Arianta arbustorum (L.) (Wiszniowska et al. 1996, Pakiet 1999). Palaeoecological analysis of the malacofauna indicates the presence of a forest with a structure resembling that of a rich Sudetic beechwood, rich herb-layer and considerable humidity, and some insolated epilithic swards (occurrence of Vallonia costata (MÜLLER)) with herbaceous plants. Palaeogeographic analysis of assemblage IV (numerous southand central-European forms, e.g. Ena montana, Oxychilus depressus, Arianta arbustorum) indicates that the dripstone cover forming the roof of the profile comes from a warm period (Atlantic), which agrees with Bosák’s (1989) conclusion on the Atlantic age of a similar dripstone in Niedźwiedzia Cave. A distinct cooling of the climate is visible in layer W2 – appearance of Discus ruderatus, regarded as typical for the lower and middle Holocene (Pakiet 1999; Wiszniowska et al. 1996) – deposited in the Subboreal period. Subsequent warming is documented in layer W1, dated as Subatlantic (more abundant occurrence of Discus rotundatus – a snail common in the area at present). The environment in the vicinity of the cave was similar in layers W2 and W1, in spite of the presence of more numerous species of cooler climate, e.g. Vitrea subrimata (REINH.), Aegopinella pura (ALD.) and Cochlodina laminata (MONT.). Climatic changes in the Subboreal and Subatlantic periods are confirmed by subfossil bat fauna, found on the surface and in the superficial part of Holocene deposits in the lower and middle horizons of the cave (Bosák 1989; Kozłowski 1989). The abundant occurrence of Myotis mystacinus indicates a slightly cooler climatic phase which is also confirmed by the presence of Myotis nattereri and Plecotus auritus, with a small proportion of Myotis bechsteini which is a forest species typical for warmer periods (Bosák 1989, Wiszniowska 1989, Wiszniowska et al. 1996; Pakiet 1999). The profile below Miniaturka Cave indicates transport of deposits from the depths of the cave system, from its still unknown parts. It should be concluded that natural zones near the entrances are unknown. The lower horizon of Niedźwiedzia Cave is located c. 35 m below the middle horizon. Till now its fragments play a role in draining the whole system (cave streams). At the same time the horizon is situated c. 10 m below the bottom of the Kleśnica stream. Consequently the conditions favour drainage of the stream waters into the karst system (periodical disappearance of flow in the Kleśnica). Deposits of the lower horizon of the cave are completely different from those of the middle horizon. Allogenic deposits, brought by the streams, prevail there. The deposits contain gravels and boulders analogous to those found in the bed of the Kleśnica. To date only scarce bone remains, mostly of bats, were found there. The age of the remains is estimated as post-glacial (Kozłowski 1989). In some parts of the lower horizons bone remains of Pleistocene animals were found, redeposited from the middle horizon. Chimneys extend from the lower horizon upwards, and some of them reach nearly to the surface (thus higher than the middle horizon). An entrance to a horizontal corridor partly Deposits and fauna of the Sudetic caves – the state of research filled by a series of at least partly cemented gravel was discovered in one of them – Komin Maurycego (Maurycy’s Chimney). The gravel rests c. 40 m above the lower horizon and is older than the dripstones of the cave which Głazek (1986) dated as 180 000 years (see Hercman et al. 1995). Regretfully the vertical position of these gravels is not determined precisely. There are indications that they may be located higher than the floor of Sala Pałacowa (Palace Hall) in the middle horizon, where the age of the bone debris is estimated as up to 40 000 years. If this were confirmed by exact measurements, the existing division of the system into horizons should be revised. Fabjańska (1987) measured the rate of recent deposition in karstic bowls of Jaskinia Niedźwiedzia. The deposit accumulated in glass Petri dishes placed on the bottom of the bowls. In areas of small human influence the deposition rate was: in Sala Pałacowa 6,1 g/m2/year (carbonates 3,4 g/m2/year), in Korytarz Diamentowy 10,3 g/m2/year 2 (carbonates 5,1 g/m /year), and in Sala Szampańska 3,9 g/m2/year (carbonates 1,3 g/m2/year). Converted to compact deposit it is 1,4-3,7 mm/1000 years, carbonates 0,5-1,9 mm/1000 years. Estimating roughly the thickness of bottom dripstone as 0,5 m it can be said that the time necessary for their formation, assuming the recent rate of deposition, was within 135-357 kA. The above-mentioned date obtained earlier by Głazek (1986) is within this range. The existing division of Niedźwiedzia Cave into genetic horizons (Pulina 1970) was based only on general and macroscopic information. 197 It should be stressed that the concept was justified only at the initial stage of the studies. The facts known at present (including the above-mentioned) indicate a need of its revision. The basis for such a revision is partly provided by the results of studies on the profile below Miniaturka Cave (Wiszniowska et al. 1996), indicating an allogenic flow in the middle horizon on the Pleistocene/Holocene boundary. The middle horizon as defined now may thus be polygenetic, with parts of it derived from periods older than the age of the bone-bearing deposits. The dripstone covers in the floor of Sala Pałacowa have not been dated so far. Their age may be however older than that of deposits with bone detritus. No unambiguous traces of human occupation or activity of Palaeolithic man were found in Niedźwiedzia Cave, though Paluch (1970) described two worn canine teeth of the cave bear, claiming that the wear was intentional and could not result from an aberrant functioning of the jaws. Świdnicki (1980) regarded damage to one of the examined skulls as resulting from a blow with a hand holding a rather big tool. In the absence of direct evidence of human presence in Niedźwiedzia Cave it can be supposed that the animal was injured far from the cave in which it then died. There was thus a likelihood of humans appearing in the vicinity of Niedźwiedzia Cave, though they did not inhabit its parts known at present. However, it should be remembered that man as a rule inhabited only parts of the caves near entrances and these were destroyed as a result of marble exploitation in the quarry. Discussion - conclusions The development of the Sudetic karst is associated with relatively small intercalations of carbonate rocks among non-karstifying rocks. Hence the cave systems occupy a much limited space. The cave horizons resulted from drainage of stream waters from valley bottoms. The altitudinal differentiation of the horizons resulted from changes in the altitude of the valley bottoms. Formation of flowing water deposits was associated with formation or transformations of the cave horizons. In later stages they could be redeposited or removed, or else covered by autogenic deposits (e.g. cave loams, dripstone layers). The bone remains are primarily associated with autogenic deposits. Redeposition is an especially widespread phenomenon in the Sudetic caves. It is 198 Jerzy Bieroński, Paweł Socha, Krzysztof Stefaniak sometimes associated with gravitational subsidence processes, including those resulting from suffusion damage to deeper horizons or water flow on the surface of the deposits. Such processes are especially important when they involve bone-bearing deposits. Redeposited bones may then suggest erroneous interpretation of the age of inorganic deposit on the basis of the bone age. The phenomenon was ignored by the pre-war researchers of the Sudetic karst. They dated their deposits based on bone remains as the period from the Tertiary till the Holocene. Reinterpretation of their results is very difficult because of the destruction of some of the described caves (e.g. Rogóżka Cave, numerous caves in Mt. Połom) or complete removal of the studied deposits (e.g. Radochowska Cave). In many cases the evidence material has been lost or destroyed. The first stage of post-war studies included an inventory and summary of the previous results of studies on the caves of the area (Kowalski 1954). Subsequently, Pulina (1977) attempted a description of the origin and development of the Sudetic karst, and introduced new concepts. One of his ideas was the concept of three horizons of the caves associated with the development of the Sudetes relief starting with the Palaeogene. At the same time he noticed the significance of redeposition for interpretation of the data. Later he initiated interdisciplinary studies on the Sudetic karst. The discovery of Niedźwiedzia Cave in Kletno was especially important for the intensification of interdisciplinary studies on caves and their deposits, organised by Pulina and Wiszniowska. The cave and its deposits were preserved in their natural state which offered a possibility of complex studies on the karst system. The results encouraged attempts at reinterpretation of earlier studies on the Sudetic caves (Radochowska Cave, Solna Jama) and at studies on the newly discovered karst localities (Na Ścianie Cave, rock shelters on Mt. Miłek). The results indicate a regional diversity of the Sudetic karst. Generally two regions can be distinguished: the western-Sudetic region (mainly Kaczawskie Mts) and the easternSudetic region (mainly the region of Kłodzko). The western-Sudetic region harbours the oldest known caves and deposits in Sudetes. The only representative of the fossil Tertiary fauna (Baranomys langenhani, probably B. loczyi) is known from that area (Południowa Cave). The eastern-Sudetic caves contain mostly upper Quaternary deposits (there is no record of older deposits). Only suggestions of a probably Pliocene age of deposits in caves on Mt. Krzyżnik were made (Pulina 1977). In the western-Sudetic region a part of the caves were within the range of Scandinavian glaciation (fluvioglacial flow). In the easternSudetic region the glaciation included only the range of Krowiarki, where however there are no larger caves. Some of the cave systems of the western-Sudetic karst (e.g. Mt. Połom and Mt. Miłek) are developed in crystalline limestones which include whole hills up to their summits. As a result several horizons of caves were formed there, correlated by Pulina (1977) with the Tertiary planation levels, with Mt. Połom and Mt. Miłek as examples. In the eastern-Sudetic karst the cave horizons are not precisely correlated with planations and their existence was suggested only in the case of Mt. Krzyżnik (Pulina 1977). References Arndt W. 1921. Beitrag zur Kenntnis der Höhlenfauna. Ergebnis einer faunistischen Untersuchung der Höhlen Schlesiens. Zoologischer Anzeiger 52, 12/13. Arndt W. 1923. Speläobiologische Untersuchungen in Schlesien. Speläologische Jahrbuch 4. Bieroński J. 1997. Hydrologia zlewni górnej Kleśnicy. (Praca doktorska). Archiwum Uniwersytetu Wrocławskiego. Deposits and fauna of the Sudetic caves – the state of research Bieroński J., Burdukiewicz J.M., Wiszniowska T. 1985. Wyniki nowych badań Jaskini Radochowskiej. Śląskie Sprawozdania Archeologiczne 26, 5-18. Bieroński J., Wiszniowska T. 1994. Zjawiska krasowe w Rogóżce (Masyw Śnieżnika, Sudety). Acta Universitatis Wratislaviensis 1072 A, 5-17. Bosák P. 1989. Osady czwartorzędowe jaskini. [in:] A. Jahn, S. Kozłowski, T. Wiszniowska (Eds.) Jaskinia Niedźwiedzia w Kletnie. Badania i udostępnianie. Ossolineum, Wrocław–Warszawa, 241254. Buczyński S., Rzońca B. 2007. Studies on the waters in the surroundings of the cave Jaskinia Radochowska [in:] P. Socha, K. Stefaniak, A. Tyc (Eds.) Karst and Cryokarst. 25th Speleological School, 8th GLACKIPR Symposium. Sosnowiec– Wrocław, Poland, March 19-26, 2007, Guidebook & Abstracts, 53-57. Croitor R., Stefaniak K. (in press). Pliocene deer of Eastern Europe. Palaeontographica. Czyżewska, T. 1989. Parzystokopytne Artiodactyla. [in:] K. Kowalski (ed). Historia i ewolucja lądowej fauny Polski, 209-217. Folia Quaternaria 59-60. Kraków. Fabjańska A., 1987. Środowisko sedymentacyjne mis martwicowych Jaskini Niedźwiedziej w Kletnie. (maszynopis) Praca magisterska, Zakład Geomorfologii Instytutu Geograficznego Uniwersytetu Wrocławskiego. Wrocław. Frenzel J. 1936. Knochenfunde in der Reyersdorfer Tropsteinhöhle. Beiträge zur Biologie des Glatzer Schneeberges 2, 121134. Frenzel J. 1937a. Die Apterygotenfauna des Glatzer Schneeberges. Beiträge zur Biologie des Glatzer Schneeberges 3, 249321. Frenzel J. 1937b. Die Reyersdorfer Tropsteinhöhle, ein schlesisches Naturdenkmal. Schlesische Heimat 1. Głazek J. 1986. Wyniki datowań nacieków jaskiniowych z terenu Polski metodą Th230U/234U. Zeszyty Naukowe Politechniki Śląskiej Mat.-Fiz. 47, Geochronometria, 2, 55-65. 199 Głazek J., Szynkiewicz A. 1987. Stratygrafia młodotrzeciorzędowych i staroczwartorzędowych osadów krasowych oraz ich znaczenie paleogeograficzne. [in:] A. Jahn i S. Dyjor (eds.) Problemy młodszego neogenu i eoplejstocenu w Polsce, 113-129. Ossolineum. Wrocław, Warszawa. Hercman H., Lauritzen S.E., Głazek J. 1995. Uranium-Series Dating of Speleothems from Niedzwiedzia and Radochowska Caves, Sudetes (Poland). Theoretical and Applied Karstology 8, 37-48. Heller F. 1937. Revision einer fossilien Fauna aus der Kitzelberghöhle bei Kauffung. Zentralblatt für Mineralogische, Geologische, Paläontologische Abteilung. A, 241-249. Kos B. 1978. Środowisko przyrodnicze Jaskini Radochowskiej oraz problem jej ochrony. [manuscript of the Msc. Thesis, Zakład Geomorfologii Instytutu Geograficznego Uniwersytetu Wrocławskiego]. Wrocław. Kowalski K. 1954. Jaskinie Polski. T. 3. PWN. Warszawa. 192 pp. Kowalski K. (Ed.) 1989. Historia i ewolucja lądowej fauny Polski. Folia Quaternaria, 59-60. Kowalski K. 1990. Stratigraphy of Neogene Mammals of Poland. In E. H. Lindsay et. al. (eds.) European Neogene Mammal Chronology, 193-209. Plenum Press, New York. Kozłowski S. 1989. Budowa geologiczna otoczenia jaskini. [in:] A. Jahn, S. Kozłowski, T. Wiszniowska (Eds.) Jaskinia Niedźwiedzia w Kletnie. Badania i udostępnianie. Ossolineum, Wrocław– Warszawa, 80-118. Leroi-Gourhan A. 1966. Religie prehistoryczne. PWN, Warszawa. Nadachowski A. 1989. Gryzonie - Rodentia. [in:] K. Kowalski (ed.). Historia i ewolucja lądowej fauny Polski, 151-176. Folia Quaternaria, 59-60. Kraków. Pakiet M. 1999. Nowe dane o holoceńskiej malakofaunie Sudetów. Geologia 25, 4, 339-362. 200 Jerzy Bieroński, Paweł Socha, Krzysztof Stefaniak Paluch R. 1970. Ślady pobytu człowieka w Seidel J. 1927. Zur Kenntnis schlesischer Jaskini Niedźwiedziej. Acta Universitatis Fledermäuse. Abhandlungen der Naturforschungen Gesellschaft. H. 1, 30. Wratislaviensis 127, 71-77. Pax F. 1936a. Die Reyersdorfer Skalski A.W. 1970. Podziemne kiełże w Tropsteinhöhle und ihre Tierbevölkerung. Polsce (Crustacea, Amphipoda, Mitteilungen über Höhlenund Gammaridae). Acta Hydrobiologica 12, 4, 431-437. Karstforschung 3, 97-122. Pax F. 1936b. Zoologische Ergebnisse neuerer Skalski A.W. 1976. Groundwater inhabitants Höhlenforschungen. Forschungen und in Poland. International Journal of Fortschritte 12, 9. Speleology 8, 217-228. Pax F. 1937. Die Höhlenfauna des Glatzer Skalski A. 1994. Fauna wód podziemnych Schneeberges. 10. Wandlungen des Polski. Przegląd Zoologiczny 38, 1-2, 35Tierlebens in der Wolmsdorfer 50. Tropfsteinhöhle. Beiträge zur Biologie des Stach J. 1936. Eine neue Art von Oncopodura (Collembola) aus der Reyersdorfer Höhle in Glatzer Schneeberges 3, 289-293. Deutsch-Schlesien. Mitteilungen über Pax F., Maschke K. 1935. Höhlenfauna des Glatzer Schneeberges. Die rezente Höhlen- und Karstforschung 3, 130-136. Metazoenfauna. Beiträge zur Biologie des Stach J. 1947. Onychiurus schoetti (LIE Glatzer Schneeberges 1, 4-72. PETERSEN), a relikt from the cave Pomorski J. 1987. A redescription of Radochów (Silesia) and its relation to the Onychiurus (Oligaphorura) schoeti (LIE group of Onychiurus groenlandicus PETERSEN, 1896) STACH, 1947, from the (TULLB) and related species. Prace cave „Na Ścianie” (Massif of Śnieżnik, Muzeum Przyrodniczego 7, 1-20. Polish Sudetes). Polskie Pismo Stammer H.J. 1936. Die Wasserfauna der Entomologiczne 57, 695-699. Reyersdorferhöhlen. Beiträge zur Biologie Pomorski R.J. 1990. Onychiurus paxi Stach, 1939, a des Glatzer Schneeberges 3, 199-214. junior synonym of Onychiurus (Onychiurus)Stefaniak K. 1995. Late Pliocene Cervids from denisi, Stach, 1934 (Collembola). Polskie Pismo Węże 2 in southern Poland. Acta Entomologiczne 60, 59-63. Paleontologica Polonica. 40, 327 - 340. Pomorski R.J. 1992. Collembola of caves and Stefaniak K. 2004. Paleoekologia i some adits of the Polish Sudetes. Acta rozmieszczenie jeleniowatych (Cervidae, Universitatis Wratislaviensis 1539, (25), Mammalia) w trzeciorzędzie i starszym 27-44. czwartorzędzie Polski. [in:] Zapis Pulina M. 1970. Wstępne wyniki badań nad paleontologiczny jako wskaźnik środowiskiem geograficznym Jaskini paleośrodowisk. XIX Konferencja Niedźwiedziej. Acta Universitatis Naukowa Paleobiologów i Biostratygrafów Wratislaviensis 127, Studia Geograficzne PTG. Wrocław 16-18 września 2004. XIV, 5-37. Instytut Nauk Geologicznych UWr. 57-59. Pulina M. 1977. Zjawiska krasowe w Sudetach Szarejko Z. 1979. Charakterystyka osadów polskich. Dokumentacja Geograficzna 2-3, Jaskini Niedźwiedziej w Kletnie. 118 pp. [manuscript of the Msc. Thesis, Zakład Pulina M. 1989. Historia odkrycia i badań. Geomorfologii Instytutu Geografii i [in:] A. Jahn, S. Kozłowski, T. Rozwoju Regionalnego UWr.], Wrocław. Wiszniowska (Eds.) Jaskinia Niedźwiedzia Świdnicki J. 1980. Zmiany pourazowe na w Kletnie. Badania i udostępnianie. kościach Ursus spelaeus ROSENMÜLLER w Jaskini Niedźwiedziej w Kletnie k. Stronia Ossolineum, Wrocław–Warszawa, 11-20. Pulina M. (Ed.) 1996. Jaskinie Sudetów. Śląskiego. [manuscript of the Msc. Thesis, Polskie Towarzystwo Nauk o Ziemi. 202 Zakład Paleozoologii, Instytut Zoologiczny, Uniwersytet Wrocławski], Wrocław. pp. Deposits and fauna of the Sudetic caves – the state of research Walczak W. 1956. Największa jaskinia Sudetów. Wszechświat 6, 133-135. Walczak W. 1958. Krasowe jaskinie Sudetów Kłodzkich. Czasopismo Geograficzne 29, 49-66. Wiszniowska T. 1967. Nowe znalezisko paleontologiczne w Sudetach. Przegląd Zoologiczny 11, 430-433. Wiszniowska T. 1970. Wstępne wyniki badań fauny kopalnej w Jaskini Niedźwiedziej. Acta Universitatis Wratislaviensis 127, 4570. Wiszniowska T. 1976. Niedźwiedź jaskiniowy z Kletna i innych jaskiń Polski. Acta Universitatis Wratislaviensis 311, 1-75. Wiszniowska T. 1978. Panthera spelaea (Goldfuss) z Jaskini Niedźwiedziej w Kletnie. Acta Universitatis Wratislaviensis 329, 113-141. Wiszniowska T. 1986. Szczątki fauny w namuliskach jaskiń na Śląsku. [in:] Dawna fauna Śląska w świetle badań archeozoologicznych. Wrocław. Ossolineum, Wydawnictwo PAN, Prace Komisji Archeologicznej 3, 9-19. Wiszniowska T. 1989. Kopalne szczątki zwierzęce. [in:] A. Jahn, S. Kozłowski, T. Wiszniowska (Eds.) Jaskinia Niedźwiedzia w Kletnie. Badania i udostępnianie. Ossolineum, Wrocław–Warszawa, 255279. Wiszniowska T., Kuryszko J. 1998. Analiza mikromorfologiczna szczątków kostnych 201 niedźwiedzia jaskiniowego (Ursus spelaeus Rosenmüller, 1784). Zeszyty Naukowe Akademii Rolniczej we Wrocławiu, 42-48. Wrocław. Wiszniowska T., Bieroński J., Pakiet M. 1996. Paleoekologia Masywu Śnieżnika [in:] A. Jahn, S. Kozłowski, M. Pulina (Eds.) Masyw Śnieżnika. Zmiany w środowisku przyrodniczym. PAE, 49-55. Warszawa. Wolsan M. 1989. Drapieżne – Carnivora. [in:] K. Kowalski (Ed.). Historia i ewolucja lądowej fauny Polski. Folia Quaternaria 59-60, 177-196. Kraków. Wołoszyn B.W. 1968. Badania nietoperzy Dolnego Śląska. Przegląd Zoologiczny 12, 208-220. Wysoczański-Minkowicz T. 1969. Próba oznaczania wieku bezwzględnego kości kopalnych metodą fluoro-chloro-apatytową. Studia Geologica Polonica 28. Zięba S., 1978. Zdjęcie morfologiczne Sal Pałacowych w Jaskini Niedźwiedziej w Kletnie. Acta Universitatis Wratislaviensis 311, 53-68. Zotz, L. F. 1937a. Die Schlesischen Höhlen und ihre eiszeitlichen Bewohner. Wilhelm Gottlieb Korn Verlag, Breslau. 38 pp. Zotz, L. F. 1937b. Altsteinzeitlicher Bärenkult in den Sudeten. Altschlesische Blätter 12, 1/2 Zotz, L. F. 1939. Die Altsteinzeit in Niederschlesien. Kabitsch Verlag, Leipzig. 143 pp.