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Excursion Guide “Cave Ecology – Terrain Course” Faculty of the Environment Jan Evangelista Purkyně University Developed by: M. HOLEC, R. POKORNÝ 2010

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Page 1: Excursion Guide “Cave Ecology – Terrain Course”fzp.ujep.cz/~Pokornyr/01_Materialy/Cave_Ecology.pdf · Excursion Guide “Cave Ecology – Terrain Course” ... This chapter

Excursion Guide

“Cave Ecology – Terrain Course”

Faculty of the Environment Jan Evangelista Purkyně University

Developed by: M. HOLEC, R. POKORNÝ 2010

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Jan Evangelista Purkyne University Faculty of the Environment Králova výšina 13 Czech Republic 400 96 corresponding author: [email protected] Foto on the introductory page: View of the stream of warm air (ventarol) from the heart of Borečský Hill. Photo: SCHKO České středohoří (Administration of the Protected Landscape Area of the České středohoří mts) Archive.

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OBSAH

Introduction ................................................................................................................................ 4

Acknowledgements .................................................................................................................... 5

What is a Cave?.......................................................................................................................... 6

Divisions of Pseudokarst Caves ................................................................................................. 7

Cave Microclimates and their Specifics..................................................................................... 9

Animal Adaptation to the Soil and the Cave Environment ...................................................... 11

The Classification of Cave Animals......................................................................................... 12

Topographic Speleology of Neovolcanites in the Ústecký kraj Region .................................. 15

Loupežnická jeskyně cave..................................................................................................... 16

Cave in Divoká rokle gorge.................................................................................................. 20

Cave on Buková hora hill..................................................................................................... 23

Literature .................................................................................................................................. 25

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Introduction The authors have been paying considerable attention to the caves in the North Bohemian neovolcanites for several years. The issue includes not only mapping the caves in the field, but also the acquisition of other more detailed characteristics. The activation of caves by invertebrates is especially studied. In the region, however, data on the occurrence of vertebrates, esp. bats, are available as well. This was the background for the idea of a school course which would call attention to the general scientific value of less known underground objects and at the same time would allow for some aspects of ecology to be studied in field conditions which for the students, despite the extensive theoretical knowledge their studies are often focused on, is rather rare. The nature of the text is more descriptive, i.e. readers are especially acquainted with geology, geomorphology, objects of fauna and the need for their protection. The functional approach to ecology, so necessary in this subject, is rather suppressed here. They are more like source materials for the study of ecology than a purely environmental text. The course is called “ecology” for simplification. We would also like to point out that our aim is not to disseminate principles of a certain subject, e.g. ecology or geology etc., but rather understanding of the studied objects. The reason is also, of course, the extent to which the course has been designed for the field, and partly the limited knowledge of relationships between individual cave organisms. The students are supposed to have a general knowledge of ecology and terminology. The aim field expeditions will be, in the examples of caves, to become acquainted with the methodologies of description of the cave environment, with individual species, especially of animals that dwell in this environment and with the methods of how to find them in nature. The choice of caves is limited partly by the conditions of state nature protection specified by the Act No. 114/1992 Coll., and partly by the course safety. In this regard, the cave Loupežnická mentioned in this guide is especially problematic. On this account, during the course only the entrance to the cave will be visited where no disturbance of bats occurs and also the risk of injury due to the difficult environment is low. The mentioned cave was, however, inserted in the list as the longest structure of a real cave-like nature. In addition to the caves presented in the text, we are always planning a guided excursion to a further and larger cave. This will provide practical experience for possible discussions about differences between large and attractive small little-known cave structures with which we are mainly concerned in this course. The course is devoted mainly to foreign students, especially to those who are engaged in the environmental field in general or in quite different specialisations from speleology or at least ecology. The aim of these texts will be as a reminder of the structures visited, especially for students. The more detailed descriptive text shows that, rather than being the basis for examination, because for foreign students it is of course absolutely unimportant to know the height of individual caves, a lot can be said at certain levels of concern and knowledge even about small cave structures, which are often not known to public. The texts should be supplemented by other chapters in the future. In particular, the part intended for students with methods of measuring abiotic characteristics in the field and methods used for sampling and assessment of animals is planned. In view of the specialisation of the authors, the course is devoted especially to animals, and not to microorganisms or lower plants that also play in these structures some, although not so much of an outstanding, role.

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Acknowledgements With regard to the fact that this material is mainly based on the book by Pokorný and Holec (2009), we would like to thank here all those who participated already on the creation of the cited book. Considering there were many people who helped and the specialisation of the course we do not mention all the names here. We would also like to thank our workplace, which allows us to explore this issue, especially within teaching activities. The quality of teaching will be also improved by the material equipment purchased thanks to a donation from the Ministry of Environment R&D (MŽP VaV) No. Sp/2d3/4/07 for the purpose of research in the years 2007–2009, and the later approved by FRVŠ project No. 305/2010/F4/a - grant of the Ministry of Education, Youth and Sports (MEYS, MŠMT in Czech).

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What is a Cave? The term cave is defined in various ways. Generally, it refers mostly to a naturally formed underground cavity accessible for people. It is obvious that from the point of view of understanding what happens underground it is not very important whether or how the man fits into the cavity or does not. Using a higher amount of definitions, we tried to set up the definition which corresponds with the orientation of our work to natural underground structures in north-west Bohemia. “Caves are cavities in the geological environment situated under the surrounding terrain level, parallel with it, or exposed at any height of the corresponding rock body. The geological environment is a part of the lithosphere formed by bodies of igneous, sedimentary and metamorphic rocks in various stages of strengthening. The cavity represents the body formed by the forces of nature (endogenous, exogenous or their combination) without human interference. The cave must be sufficiently long in the major part of its length so that an average adult can walk or crawl through it. This fact does not eliminate the presence of a component of the cave or of the cave system that cannot be crawled through – components in the shape of narrow to dying-away endings, side branches, slots, etc. of any length. To fulfil the definition of a cave, the total passable length of the structure should be long enough to be able to significantly distinguish the inner environment from the surrounding environment, especially on the basis of the presence of a different microclimate, humidity, the presence of an aphotic zone, cave sediments and troglobiont or at least troglophile organisms. In practice, this limit was empirically set to the distance of 5m from the outer environment. Caves are also natural underground spaces corresponding to the dimensional criterion, but without a passable inlet orifice, or without any connection with the outer environment, if their presence is reliably confirmed using survey methods.”

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Divisions of Pseudokarst Caves Epigene pseudokarst caves can be distinguished from the point of view of their formation and divided into several groups. They are caves of crack, cleft, talus, erosion and combined types. The specific type is a tree mould cave, which will not be discussed here. The terminology is based on studies of VÍTEK (1981, 1979) supplemented with the findings of POKORNÝ and POKORNÁ (2007) and ESZTERHÁS (2007). Although the studies of Vítek and Eszterhás are devoted mainly to sandstone, their classification is so general that it can be applied, in addition to sediments, to both igneous and metamorphic rocks. This chapter describes mainly those genetic cave types representatives of which can be found in the neovolcanites of the district Ústecký kraj. The first two types, crack and cleft caves, are genetically connected to the movement of rock blocks and according to ESZTERHÁS (2007) therefore belong to one category. Crack caves represent vertical narrow spaces where the height significantly prevails over the width and at the same time they pass the required criterion of necessary length. Characteristic signs are a trapezoidal cross-section and a ceiling formed by rock material connected with upper layers of rock. Their formation is always predisposed by the existence of a system of more or less vertical concurrently converging cracks – zones of substantial tectonic disruption. The cave is identified as the crack type on the condition that the rock mass affected with the fan of tectonic failures (cracks) remains stable as a whole. If this condition is not met, the cave is categorized as the cleft type. In these cases block movements of rock blocks up or down the slope occur and underground spaces are gradually widened and opened as a result. The rock can be cracked in the same way as in the previous case, however generally the steep

relief of the landscape will start the process of loosening individual blocks along

tectonic predispositions and their sliding to lower levels. The space formed is usually narrow and high in cross profile – according to the method of formation – in

the classical form either roof-shaped, in the shape of letter “A”; or wedge-shaped, in the shape of letter “V”. In the second case mentioned the ceiling is formed by accumulated and embedded rock blocks. The classic example of a crack cave is Bílanina. Examples of cleft caves in the Ústecký Kraj region are the Komora cave in the Lužické Mountains or Loupežnická Jeskyně (the Robber's Cave) near the town of Ústí nad Labem. The caves whose formation is connected only with cambering along the planes of division of rockslides are according to PANOŠ (2001) called rock-slide caves.

Fig.1: A crack cave. Illustrations from the authors’ archive.

Fig. 2: A cleft cave of the “A” type. Illustrations from the authors’ archive.

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The North Bohemian Region is quite rich in geomorphological objects during the formation of which frost weathering played a role in the Quaternary period (RAŠKA and CAJZ 2008). Various rock and boulder streams and also rock cabins, frost cabins and non-karst caves can be found here. The origin of these “frost caves”, whose classification is difficult, is connected with the intrusion of ice and the opening of cracks. Consequently, the shift of rock bodies in these cases is not influenced by a steep slope. Considering the fact that their genesis is connected with at least the partial movement of rock bodies, these caves are classified in one category of the cleft type although for some of them the classification of frost caves could be used as well, as PANOŠ (2001) states in his survey. Such cryoplanation cleft caves can be found in neovolcanites of the Ústecký Okres region on the Buková Mountain (e.g. the cave Ledová Jeskyně) and the cave Jeskyně Skřítků. Other types are caves of the block-field type or talus type. Although these structures are not considered by some authors (e.g. OZORAY 1962) to be caves in a narrower sense of the word and the terms like pseudo-caves have been suggested, it is beyond any doubt that they are caves when all the criteria such as length, if it is passable, etc. are met. The formation of talus caves is, in the same way as cryoplanation clefts, often connected with frost weathering. The

longest Czech talus caves are several hundred meters long and it is typical that they are not passable for an extensive section. The category of subsurface talus in the region is represented by, for example, the cave Jeskyňka near Dobrná and also by a smaller cave on the Buková Mountian and around the cave Jeskyně Skřítků. Other types are caves of the erosion type, which includes caves formed by water and wind erosion according to ESZTERHÁS (2007); caves during the formation of which internal stress in rocks or temperature variations played a role and also caves formed by chemical erosion (dissolution, hydration, hydrolysis,

oxidation), including bio-erosion. VÍTEK (1981, 1979) in his studies comments on bedding type caves, which are a large subset of erosion caves including structures formed by weathering and the removal of less resistant bedding planes or the crumbling away of layer banks. The formed spaces are low, relatively wide and of various lengths. According to VÍTEK (1981) bedding type caves occur only in sedimentary rocks, whereas similar objects in eruptive rocks and metamorphites according to him are crack caves based on horizontal cracks or planes of division. However, POKORNÝ and POKORNÁ (2007) identified within the Czech Central Mountains several caves and small caves corresponding to bedding type caves, e.g. in sandstone at Český Ráj. For more details see POKORNÝ and POKORNÁ (2007).

Fig. 3: A cleft cave of “V” type. Illustrations from the authors’ archive.

Fig. 4: A talus cave. Illustrations from the authors’ archive.

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Cave Microclimates and their Specifics PŘIBYL et al. (1992) define three basic types of caves in the relation to their microclimate, where the nature of the circulation of air masses in the system of inner/outer environment is considered to be the main classification criteria. The first type is a static cave. There is generally one entrance to these caves or if there are even more entrances, they must lie at the same altitude. As long as the floor flow comes down in the direction of the entry below its level, in winter season the lighter warm air begins to escape up out of the cave and at the same time the outer cooler air is sucked in. In summer the circulation stagnates, since the heavy cool air is kept inside the cave and warming in the direction of the depths from the entrance is only slow. Static caves with the floor falling from the entrance down are called cool caves. In the case of the cave cavity being situated above the entry level, a quite opposite effect occurs. In summer season the cool air spontaneously flows out of the cave and the warmed outer air is sucked in. Later, in winter season, this reservoir of warm air is cooled down by the surrounding rock only slowly and in winter, when the stagnation of circulation occurs, the temperature in these static warm caves will to be markedly higher than outside the cave cavity. Static caves are thus generally characterised by air circulation only during some defined part of the year (cool ones in winter, warm ones in summer). However, similar caves are rather rare, because most of the caves, karst or non-karst, are established in a rock mass interwoven by sets of cracks, bedding joints and slots. Small cavities generally allow passage to the surface and due to that continual circulation begins and continues for the whole year. Relatively frequent are also cases when there are multiple inputs to the cave (both passable and non-passable) at various altitudes. These caves are categorised as dynamic. The air circulation in dynamic caves proceeds according to the scheme whereby in summer the air circulates from the upper entrance in the direction of the entrance situated below, whilst in winter the opposite occurs. In summer the driving force is the cool air inside the cave which, due to its higher weight, flows out of the cave through the lower entrance and from above is substituted with the warmer air from outside. The intake air is cooled down through the heat transfer to the rock due to which the relative air humidity is increased. When the air is 100% saturated, the subsequent precipitation of humidity occurs. In contrast to static caves, the circulation cycle in winter is not interrupted, but due to the outer temperature decrease it reverses. The inner warmer thus lighter air exits through the upper orifice and the cool air is continually sucked in through the lower entrance of the cave. Along with the cooling of the rock from the cool air in winter, the air humidity in the cave when compared to

Fig. 5: Top left is the static cave with the upper entrance maintaining the cool climate during the whole year. Top right is the static cave with the lower entrance where the cool air accumulates during the whole year. Bottom left is the dynamic cave with air circulation during the whole year. Freely according to MUSCIO in STOCH (2002).

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the summer heating of the rock is decreased and is supplemented by evaporation from the walls. So it can be generally said that in dynamic caves the walls in summer are dewy and damp, and conversely, in winter these caves become significantly drier. According to PŘIBYL et al. (1992), statodynamic caves are sporadic and specific. Their basically dynamic nature is interrupted for some part of the year due to the closing of one entrance, e.g. by temporary glaciation or flooding. This place then acts as a stopper that prevents the circulation of air masses. Similar air circulation occurs also in cracked massifs or rocky accumulations. Places with warm winter flow streams were in the past incorrectly connected with volcanic activity and the improper term “fumarole” was used (see e.g. SOMMER 1833, POKORNÝ, 1931), which comes from the Latin word fumus, meaning smoke. Earlier authors speculated about the existence of a crack system extending to such depths that the air could be heated from thermal groundwater bodies extending from the foothills of the Ore Mountains (e.g. ŠIMR 1957). Only later was it found (specifically by VÁNĚ 1992) that they are manifestations of air circulation and therefore the term ventarols (ventus = wind), which better represents the principle of the phenomenon, was suggested. Places where cool air and even snow and ice remain till late spring are called ice pits. Provided that it is a cave, the term ice (where ice remains in the cave for the whole year) or pseudo-ice cave is used, according to the time for which ice remains in the cave. According to HROMAS (1971) there are no real ice caves in the Czech Republic. The various “ice caves” are therefore pseudo-ice caves. Pseudo-ice caves are definitely more frequent . With the character and nature of the climate they are very similar to ice caves, but due to a coincidence of factors (small size and length, partial insolation of the entrance, periodic through-flow of water, etc.) the ice is present in these structures only until spring or, at most, until summer. For some part of the year, these caves are completely without ice. For pseudo-ice caves thick snowdrifts of firn snow in the cave portal are characteristic. The snowdrifts are reservoirs of snow and at the same time they maintain the cool air inside the cave. Among the karst pseudo-ice caves in the Czech Republic that can be named are Ledová Jeskyně cave in the Moravian Karst or Blátivá Chodba cave in the Chýnov caves. In the category of non-karst caves, Ledové Sluje (Ice Caves) near the town of Vranov nad Dyjí should be mentioned. It is probably the longest cave in the Bohemia, well explored by professionals, with the total length of up to 400m (HROMAS and BÍLKOVÁ 1998). About 10m long, Ledová Jeskyně cave in the body of Velký Bezděz is only briefly described by CHVÁTAL (1996) and HROMAS (1971) and is substantially less well-known. In the Ústecký Kraj region there are also structures with the character of pseudo-ice caves. In particular, there are cleft caves on the Buková Mountain, some of which have their cool nature already in their name – Ledová Jeskyně (Ice Cave) and Sněhová Jeskyně (Snow Cave). The former has already been mentioned in literature almost two hundred years ago, when (SOMMER 1833) spoke of the “30 feet deep crack where snow can be found even in late summer”. PLEISCHL (1838) published the discovery of several deep clefts near the Buková Mountain and stated that during his visit at the end of August there was already no snow. ZAHÁLKA (1890) describes in the vicinity of the peak of the Buková Mountain the 12m long cavity with a North-South orientation where an icy glaze forms even in the early summer months. It can be found in travel literature that in the vicinity of the cave Ledová Jeskyně cryophilic species of plants grow due to the long presence of ice and snow, e.g. Rosa pendulina (alpine rose) and Stachys alpinus (stachys alpina) (BREDSCHNEIDER 1928). The presence of cool air to a certain extent simulates the character of the northern climate which is confirmed by the presence of various cryophilic organisms – e.g. the mite Rhagidia gelida (ZACHARDA et al. 2005) and also the northern European spider Semljicola faustus

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(ČEŘOVSKÝ and HOLEC 1996). Conversely, the existence of ventarols and thus places where the temperature even in winter doesn't drop below freezing enables on the Borečský Hill the occurrence of the thermophile Mediterranean liverwort Targionia hypophylla, only at this one locality in the Czech Republic (KUBÁT 1971). Probably the best known places where ventarols and ice pits occur are the national nature monument Borečský Vrch (esp. ventarols), the nature monument Plešivec in the district of Litoměřice (esp. ice pits) and also the nature reserve of Kamenná Hůra east of Děčín (well developed ice pits and ventarols). These structures are so significant that their conservation is protected by the declaration of small-scale conservation areas where the main object of conservation is the occurrence of ventarols and ice pits.

Animal Adaptation to the Soil and the Cave Environm ent According to the distance from the ground’s surface there are three main types of environment for organisms: epigeic – on the soil surface, endogeic – the inner environment of the soil, and hypogeic – the environment under the soil surface, e.g. the cave environment. The common property of soil and caves when compared to open land is a lack or even absence of light and a more stable climate. For many soil (especially endogeic but sometimes also epigeic) and cave species the loss of body pigment, the total reduction of the eyes, the reduction of wings, etc. can be observed. These characteristics are very remarkable and can intuitively be best explained relative to the adaptation of these species to the dark environment. In the case of cave species we speak about troglomorphism, in the case of soil species about euedamorphism, which is the less frequently used term nowadays (see e.g. DUCARME 2004). The above mentioned characteristics used to be called sensu stricto troglomorphisms, in contrast to larger sense, thus analogically sensu lato troglomorphisms, where, when defining troglobionts and troglomorphisms, also other (physiological, ethological, reproductive) responses of organism to the environment are considered (e.g. CHRISTIANSEN 1992, 1962). In addition, for etymological reasons, the term troglomorphy should be substituted by the term troglobiomorphy according to JUBERTHIE and DECU (1994); many authors respect this, although other authors use the more traditionally established term. Movement in small soil spaces is limited by the development of various body appendages. E.g. a furcula (a little fork-like springing mechanism) so typical for epigeic species of collembolans is strongly reduced in endogeic collembolans, because it’s very difficult to jump in soil. The situation is similar with other appendages, especially limbs, antennas, etc. Conversely, in caves, or at least on the surface of caves and not in their sediments, which are also inhabited, elongation of limbs and antennas, etc. are more likely be

Fig. 6: Example of the reduction of the eyes at the spiders Porrhomma egeria. The independent reduction of individual eyes and variability of the reduction are interesting. Simplified according to SANOCKA (1998).

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observed. The reason is that in the dark environment with a small amount of food it’s necessary to have long appendages acting as elongated senses. The creature cannot see in the dark, but thanks to the elongated part of its body, it can feel its surroundings. The comparison comes to the mind of a man groping in the dark with his hands outstretched, or a blind man with a white cane. In addition to the different amount of space offered by soil and caves, there are other differences. Especially the availability of food, when compared to soils in caves, is generally very low. The main difference between cave animals and non-cave animals may in particular be the lack of ability to compete with organisms from the outside world (SKET 2008). Among other adaptations of cave organisms are slow movement and low reproductive ability. Both the reluctance of species from the outer environment to penetrate into caves and the reluctance of cave species to leave, i.e. to enter an environment with higher food availability but at the same time more competitive species, can be explained by the low food availability in caves (SKET 2008).

The Classification of Cave Animals The goal of various classifications is to organise knowledge into a well-arranged and practically usable system. Often it is a simplification at the expense of precision. In practice, then, we can either use more simple, but sometimes less precise classification, or more detailed classification, but which often demands more information about the classified object. The situation is not different for the classification of cave organisms. For the classification of cave, or cavernicolous (Latin caverna=cavern, colere=to inhabit) species, the simplest approach seems to be to divide them into cave and non-cave organisms, or into those that cannot be unambiguously classified as one of these two categories. Traditionally troglobionts, troglophiles and trogloxenes are distinguished. With regards to the often less well-known bionomics of various species and a variety of classifications where the meanings of the same terms overlap or even new terms are used, the use of classifications is rather problematic. In contemporary literature, the exact meaning of the term according to the sense in which the author meant it or in which it is used should always be mentioned. Troglobionts (Greek trogle=cave, bioteuo=dwelling) Troglobionts are real cave animals, or more simply – just cave animals. The presence of troglomorphisms plays an important role in the definition of troglobionts in speleobiological literature (other specific adaptations are often unknown for particular species). CHRISTIANSEN (1962) defined: a. troglomorphic taxa, i.e. highly modified forms when compared to epigeic taxa, b. ambimorphic taxa, partly modified and similar to epigeic taxa, c. epimorphic taxa, i.e. dwelling and reproducing in the cave, but without morphological modifications, d. trogloxenous taxa, i.e. taxa in random caves or exploiting caves, but without specific adaptations. In our conditions, morphological adaptations as a criterion for the delimitation of species of spiders dwelling in underground environment of stone runs were used by RŮŽIČKA (2001). Some species, however, do not manifest characteristic morphological properties, or these properties are manifested also by species living outside caves (see e.g. CULVER 1982,

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MATILE 1970). It is not important if the biotopes are “similar” (e.g. in the sense of similar humidity, temperature, dynamics of environment changes, etc.) or not. For example SKET (2008) reports the example of the freshwater crustaceans (shrimps) Niphargus where the reduction of the eyes is not dependent on life under the ground. Differences can also be found in animal behaviour, their physiological processes, etc. On this account, a purely morphological approach is rejected by some authors for the definition of real cave animals. Whatever the criterion selected, i.e. whether it’s morphological, physiological or other, we always look for the properties that we expect in the organism and only on their basis do we determine whether it is a cave animal or not. This means that we tend to determine its relationship to caves indirectly. It can be true that a given species or population is permanently connected with caves, i.e. it is a real cave organism, but the time when selection operates is short and no obvious and expected adaptations occur, or that these properties are not influenced by any selection in this case. Many cave species with no known surface populations are categorised as troglophiles due to the fact that they show only indistinct signs of regressive evolution (pigmentation loss, eye reduction, etc.) (CULVER 1982). The intuitive explanation of adaptations “apparent at first sight” may, with some problematic exceptions, meet another problem. As was mentioned above, some cave species have long limbs “because” there is little light in the cave, so the food cannot be seen and must be felt for, which is probably easier with long limbs than with short ones and this fact, from the point of view of evolution, is so significant that this selection really occurs. But it is based on the general presumption of the lack of food in caves. Locally, however, there can be enough food, e.g. in the environment of bat faeces. The generally used term of an abundance of food is then also problematic. Theoretically, this adaptation should be advantageous only in dark and food-poor caves. This means that even short leg forms should be considered troglobionts. It could also be the problem of various morphometric measurements that underground species often have larger bodies. For example DUCARME et al. (2004) confirmed that the cave mites he studied generally have larger bodies and at the same time greater length of some limbs, trichobothria, etc. than soil species. After taking into account the lengths of these mentioned dimensions relative to body length, the differences between cave and soil species disappeared. The result from this approach was that only one sign remained from several possible signs. Troglophiles (Greek filéo=to love) In the sense of the classification proposed by Schinner-Racovitza this is a marginal group between entirely cave-dwelling species (troglobionts sensu SCHINNER 1854 and RACOVITZA 1907) and species dwelling in the outer environment who only occasionally visit caves (traditional trogloxenes sensu SCHINNER 1854 and RACOVITZA 1907). According to these authors, troglophiles are regularly found in caves with daylight and rarely outside them. Other classifications have been offered by many authors, among which it is important to name RUFFO (1957), PAVAN (1944), DUDICH (1932), etc. For example, according to the Italian author PAVAN (1944), troglophiles are those species that prefer the underground environment, but are not adapted to it, i.e. do not show special adaptations. Another Italian author, RUFFO (1957), defines so-called eutroglophiles, species able to live externally, but preferring to be underground and able to reproduce there as well, i.e. they are surface species able to form more or less stable populations underground. However, this is what BARR (1968) considers “troglophiles”. In the traditional system of Italian speleobiologists, in addition to eutroglophiles, the group of subtroglophiles is used. They are the species that exploit the cave environment, but the non-cave environment is used by them for at least one life function (food, reproduction etc.).

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Fig. 7: A possible schematic expression of the classification of cave animals (organisms). Trogloxenes (Greek xénos=foreign) In the sense of the traditional classification proposed by Schinner-Racovitza (Schinner-Racovitza’s system), these are animals “foreign to caves”, i.e. species that got into a cave more or less randomly. In spite of the fact that this is a group over which there is least conflict, even here we face nomenclature troubles. Whereas the above mentioned Italian authors use the term trogloxene in the sense of Schinner-Racovitza’s system, BARR (1968) did not use this term in the sense of subtroglophiles and instead of the term trogloxene used the term “accidental”. He thus introduces a category which is underground even less than trogloxenes. In principal the same classification, based on a classification according to the environment, is also the recently published classification proposed by SKET (2008) with the following categories: troglobionts, eutroglophiles, subtroglophiles and trogloxenes. With regard to variety of terminology and classification problems, the use of the chosen unified scale (e.g. the one proposed by Sket) would facilitate the orientation in the problem. Troglobionts (troglobites, eutroglobiontes, obbligato troglophiles, in water environments so-called stygobionts, etc.) should, according to the author, be the species most strongly related to the underground environment. A population or subspecies that forms a part of an eutroglophilous species can even be troglobiotic. Eutroglophiles (partial troglophiles, facultative troglophiles and hemitroglobionts) are species which are essentially surface species, but able to maintain constant underground populations that can be troglobiont. Subtroglophiles (partial troglophiles, pseudotroglobionts and trogloxenes) are inclined to live (permanently or temporarily) underground, but they use the outer environment for some biological functions (daily – food; seasonally or throughout their life – reproduction). Trogloxenes (“accidental”, eutrogloxene, tychotroglobiont) are the group of species that sporadically occur in the underground environment and are not able to form underground populations (SKET 2008).

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Topographic Speleology of Neovolcanites in the Úste cký kraj Region The Ústecký kraj region by no means belongs to areas interesting in terms of caves. The reason is the absence of the limestone rocks that enable the presence of classic karst underground scenery. That is why stalactites, stalagmites, cave pearls and other attributes of a “classic” cave cannot be found here. Nevertheless, there is no lack of underground spaces. Not only the Ore Mountains, but also the Czech Central Mountains are interwoven with literally hundreds of galleries, shafts and old mine workings, the remains of ore and brown coal mining. Rock climbers scaling walls in the Elbe valley north of Děčín know well the narrow and steep cracks between rock blocks which are often interconnected with a net of squeezes (squeezes or crawl spaces - small caves or spaces only passable with effort because of its small dimensions) caves – caves tens or hundreds of metres long in the sandstone rocks). The rocks that dominate the district Ústecký kraj – the Czech Central Mountains – are indeed of speleological interest. They are formed from Tertiary volcanites with slopes and peaks often weathered into bizarre shapes and forms. A relatively inconspicuous part of the range of peaks of the Czech Central Mountains is not very long and deep caves and small caves that are cut into the rock. So far several dozen of them have been described and, though it’s a wonder, also the longest recorded cave belongs to them. It is the cave Loupežnická jeskyně (the Bandit’s Cave) near the town Velké Březno and with its 130m length it represents the longest cave in neovolcanic rocks in the Czech Republic. Although the average length of local caves is shorter by at least one order of magnitude, on the next pages you can find the detailed description of all the structures meeting the definition of cave. We would like to introduce readers to genius loci of these hidden and inconspicuous places.

Fig. 8: Schematic cartogram of the location of cave formations of interest in the Ústecký Kraj region. 1 –Loupežnická Jeskyně (the Robber’s Cave), 2 – The niche on Magnetovec Hill, 3 – The niche on Sokolí Hřeben Mountain, 4 – The Průčelská Rokle caves (the cave Skluzavka, the small cave Medvědí Brloh), 5 – The cave in Divoká Rokle valley, 6 – The cave on Buková Mountain, 7 – The Komora cave, 8 – The small cave near Dobrná, 9 – The small abyss at Hanziho

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Propástka, 10 – The small cave Liščí Jeskyňka, Ve Stěně cave, U Cesty cave, 11 – The cave on Pustý Vrch hill, 12 – Zlatá Díra cave (Goldloch), 13 –Pod Vodopádem cave, Kobka cave, the cave near Františkov n. Ploučnicí, 14 – The small cave Valkeřická Jeskyňka, 15 – The niche in the rock Bechlejovická Stěna, 16 – The small cave Lomená Jeskyňka, Komínová Jeskyně cave, 17 – Bilanina Jeskyně cave, the small cave on Bořeň Hill, 18 – The cave and the small cave on Radobýl Hill, 19 – The small cave on Výhled Hill, 20 – Mariánská Jeskyně cave, 21 –Natrolitová Jeskyně cave, 22 – Ledové Díry cave on Kamenec Hill, 23 – Sklepení cave on Kamýk Hill, 24 – Sedlecká Chodba cave. The grey areas on the map indicate the presence of neovolcanic rocks. Loupežnická jeskyně cave (50°39,648´ N, 14°7,560´ E) Location: Loupežnická jeskyně cave (could be translated as the Bandit’s Cave), in German literature mentioned as Räuber - or also Räumerhöhle, is located 5 kmto the East from Ústí nad Labem on the northern slope of Zámecký vrch hill (elevation of 311.9 m above sea-level) (in old maps marked as Hanberg or Pfaffenberg). Geology and geomorphology: The whole cave was formed in the nefelinite trachyte rock called also phonolite. The cave at Olešnice is definitely the longest and deepest cave in neovolcanic rocks in the Czech Republic. It was probably formed by frost weathering, sliding of slopes and widening of rifts in phonolite during Quaternary. Description: The entry is 0.8 m wide and 0.4 m high, it is situated to the north. The entry is protected by a loose grid and it is freely accessible. The main rift direction is at first northeast – southwest. About 10 m from the entry there is the first rock step going straight down. Further the road becomes larger in width and height, however at the end of this space there is a vertical wall with narrow triangular manhole at the rock foot. Speleologists call this place ”Guillotine“ according to the high sharp stone that is hanging under the ceiling about 5m directly above the manhole through which one has to crawl to get into the neighbouring space. The total length of the cave which could be accessed is about 120 m, however the total length of accessible roads is much bigger. As the slopes of the valley of Elbe are not stabile in general it can’t be excluded that the Loupežnická jeskyně cave will change in shape also in future, but these would be more probably sudden changes. The cave has been protected since year 2001 as Natural sanctuary Loupežnická jeskyně cave of total area 13 hectares, it is situated in CHKO(protected landscape area) České středohoří and at the same time it is important recorded and monitored wintering for specially protected species of bats (CHVÁTAL and KINSKÝ 2000). Fauna: Loupežnická jeskyně cave is the biggest accessible natural underground space (i.e. cave) in the region of Ústí nad Labem. Due to the fact that fauna of the Czech Republic is not much developed it is likely that also in this location there are only those types we can find in some other caves of the region of Ústí nad Labem, or in other areas of the Czech Republic. Special protection is provided to Chiropterans, especially lesser horseshoe bat (Rhinolophus

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hipposideros) and large mouse-eared bat (Myotis myotis). Both species are critically endangered based on the bill of the Ministry of Environment 395/1992 Col. Other endangered species in this location is highly endangered fire salamander (Salamandra salamandra). Noticeable species of invertebrates living in this area are for example: spider Porrhomma myops, which is entirely linked with underground or rove beetle Quedius mesomelinus, which is also typical species for caves but it can live also on surface. Remarkable species of invertebrates is especially a big spider of honey colour Meta menardi. Protection of the object and its fauna: As the cave is just a nature sanctuary and it is not protected by any European procedures, which means it is not SAC locality. The entry for visitors should be free. The only exception is entry due to the research reasons. The entry should be allowed in advance by the authority of nature preservation (authority of the protected landscape area České středohoří). Due to the fact that critically endangered species the lesser horseshoe bat and the mouse-eared bat occur in this location frequent entries might disturb and endanger their populations so the frequency of visits should be also controlled and permits should be required to enter the location. However these procedures are not normally adhered to and it is practically impossible check and punish those who breach the rules. Therefore each visitor should consider the entry, i.e. people should be aware that they might endanger rare species. Especially when more people enter the cave the air can get warmer and the bats are disturbed. The excursion will go just to the entry or possibly to the front part of the cave. The excursion itself would be discussed in advance with the Authority of Protected Landscape area České středohoří (Správa CHKO České středohoří). The employees of this authority and people from AOPK ČR monitor the bats on regular basis.

Fig. 9: Entries are in some points very narrow.

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Fig. 10-12: From top to left: large mouse-eared bat, lesser horseshoe bat and fire salamander.

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Fig. 13: Scheme of Loupežnická jeskyně cave. Modified based on mapping of Mr. Vavřinec and Mr. Duben in CHVÁTAL and KINSKÝ (1996).

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Cave in Divoká rokle gorge (50°40,864´ N, 14°07,809´ E) Geology a geomorphology: On the way from Ústí nad Labem to Děčín you can’t miss the sky-line of trachyte hill called Kozí vrch (379.6 m above sea-level) on the left bank of the river of Labe opposite the urban neighborhood Mojžíř. Kozí vrch hill is located in area which was markedly influenced by tectonic activity. The results of volcanic activity could be easily observed in near Divoká rokle gorge, which is opening 500 m to the north-west from Kozí vrch hill. The interesting phenomenon of this gorge is that already during the period of the volcanic activity there was a deep depression in Mesozoic sandstones which was 2 km long, 800 m wide and 200 m deep. This depression was under the water level of a large lake. Inside the depression there were settling layers of material from volcanic eruptions. Relatively recent land slide enabled formation of small pseudokarst cave. The cave is located approximately in the middle of the height of the gorge in its left slope about 3m above the lowest place of the narrow gorge profile. Description: The cave is located approximately in the middle of the Divoké rokle gorge at Kozí vrch hill, in its left slope about 3 metres above the lowest place of the narrow gorge profile. Roughly in the middle of the length of the rock outcrop, there is an oblique and high crack where the entry into the cave formed. The shape of the gate is pointed triangle with bottom length of 130 cm and leg lengths of 220 and 240 cm. First roughly 2 m of the length of the cave are of the rift nature and the cavity is of a letter “A” shape with gradient of 45°. In the back right part of this atrium there begins a small hemispherical tunnel. After next two meters the tunnel leads into the final wider cavern. The width of the cave at this point is up to 2 m, the height is up to 1.1 m. The end of the cave is sudden, in the cavity there are not any blind drifts. The total length is 7.1 m. The bottom of the front of the cave is partially covered by leaves, in the tunnel the bottom is covered by the layer of waste and rock fragments. The first half of the cave is sloping under the angle of 30° down the cave and then the angle even increases to about 50°. Fauna: Fauna of the cave is not known in detail. However one of the most significant species is spider Meta menardi. Preservation of the object and its fauna: The cave in not subject to the protection according to any special procedures. Entry is free. Only in case of research a permit of nature preservation authority is required but it is more or less formal in this case. The cave is not suitable for wintering of bats so there is no need to worry about their disturbing. The cave can be accessed easily so any destructive activities usually done by speleologists to enlarge entry or galleries are not expected.

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Fig. 14:View of the entry part of the cave in Divoká rokle gorge.

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Fig. 15: View of the entry part of the cave in Divoká rokle gorge.

Fig. 16: Ground plan of the cave in Divoká rokle gorge.

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Cave on Buková hora hill A rounded peak of Buková hora hill (Zinkenstein, Tzienkenstein, Buchberg) (683.3 a.s.l.) is a significant dominant of the east part of České středohoří mountains. You recognize it easily because on the top of the mountain there is a television transmitter rising up to 223 m. Buková hora hill was in the past a popular tourist destination. At the turn of 19th century there was a view point with a wooden platform. This place is called Humboldtova vyhlídka (view point of Humboldt) and you can climb up the hill following a red marked footpath. Buková hora hill and its surroundings are formed by few extrusions of basaltic and tephrite rocks. Due to the relatively high altitude many rock cliffs, single small rocks and stone fields (nowdays forested) could have been formed in top parts of Buková hora hill during the Quatenary. The rocks are significantly cracked up in two upright directions northwest –southeast and southwest-northeast. Combination of landslides, tectonic predisposition and of frost weathering is probably the cause of formation of few small caves. We will mention here the most famous one called Ledová jeskyně cave. Ledová jeskyně cave (50°40,262´ N, 14°13,696´ E) Location: The cave is located on southeast slope of Buková hora hill below the edge of tableland in the altitude of about 660 m. There is a footpath going from the television transmitter down the hill into the forest. After about 30 m there is 5 m high rock step in the slope formed by tephrite, down to the right there is another 6 m high rock step. The cave starts at the bottom of the second rock step and its nature is of a vertical rift leading towards the rock massif. Description: Entry to the cave is double it is facing southwest and it is freely accessible. The cave is formed by a rift of northwest-southeast direction, which is sloping downwards, superelevation from the entry to the end of the accessible part is about - 2 m. Width of the cavity varies between 30 – 50 cm, height is 3.5 – 4.0 m. Length of accessible part is 7 metres, at the end the cave is blocked by a huge rock and some smaller stones. Fauna: Significant representatives of fauna in this location are for example troglophiles of rove beetles Quedius mesomelinus, Tachinus rufipennis and Omalium validum. Other noticeable species are Cave Spider Meta menardi or the Herald butterfly (Scoliopteryx libatrix). Preservation of the object and its fauna: The cave in not subject to the protection according to any special procedures. Entry is free. Only in case of research a permit of nature preservation authority is required. The caves on Buková hora hill allow entry of bats into deeper parts of cracked rocks therefore these are important locations from the chiropterological point of view.

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The bats can be rarely seen in the accessible parts of the cave so there is a minimum chance of their disturbing.

Fig.17: View of Buková hora hill with the television transmitter.

Fig. 18: Ground plan of Ledová jeskyně cave.

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