Faculty of the Environment Jan Evangelista Purkyně University in Ústí nad Labem Subterranean Habitats M. Holec, R. Pokorný Ústí nad Labem, 2012 Faculty of the Environment Jan Evangelista Purkyně University in Ústí nad Labem Subterranean Habitats Název: Subterranean Habitats Autoři: Mgr. Michal Holec, Ph.D. Ing. Richard Pokorný, DiS. Vědecký redaktor: Prof. Ing. Jaroslav Boháč, CSc. Recenzenti: RNDr. Miloslav Zacharda, CSc, RNDr. Karel Tajovský, CSc. © Fakulta životního prostředí Univerzita J. E. Purkyně 2012 ISBN: 978-80-7414-416-5 (brož.) ISBN: 978-80-7414-897-2 (online: pdf) Tato publikace vznikla v rámci projektu OPVK EnviMod - Modernizace výuky technických a přírodovědných oborů na UJEP se zaměřením na problematiku ochrany životního prostředí. Reg. č.: CZ.1.07/2.2.00/28.0205 Obsah Acknowledgements . 5 Introduction . 6 Emergence of Selected Underground Spaces . 7 Pseudokarst Caves . 10 Definition of Types of Natural Underground Spaces . 12 Cave ....................................................................12 Cavelet ..................................................................13 Chasm...................................................................13 Rock overhang ............................................................13 Superficial subterranean habitats ..............................................14 Stony accumulations........................................................15 Most Important Cave Regions in the Czech Republic . 16 Bohemian Karst ...........................................................16 Moravian Karst............................................................17 Branná Belt Karst ..........................................................18 Hranice Karst .............................................................18 Javoříčko Karst............................................................19 Mladeč Karst .............................................................19 Pálava Hills Karst..........................................................20 Chýnov Karst .............................................................20 Karst of the Kamenice and Železný Brod catchment basin ..........................21 Železné hory Karst ........................................................21 Tišnov Karst ..............................................................21 Main pseudokarst areas in the Czech Republic ...................................21 Subterranean habitats from the viewpoint of protection and administration . 23 Cave utilization............................................................23 Cave management and care ..................................................23 Cave research and evidence . 25 Environmental Conditions and Their Measuring . 26 Subterranean Microclimates..................................................26 Light ....................................................................28 Heat (temperature) .........................................................30 Humidity.................................................................35 Organic matter in subterranean habitats .........................................38 Atmosphere in subterranean habitats . .41 Methods of animals’ collection . 48 Direct searching for animals..................................................48 Pitfall trapping ............................................................49 Core sampling of soil, sediments or litter and fauna extraction.......................49 Ecological and Evolutionary Classifications . 51 The origin and emergence of subterranean terrestrial fauna . 55 Survey of selected groups of organisms . 57 Nematodes (Nematoda) .....................................................58 Annelids (Annelida) ........................................................59 Molluscs (Mollusca)........................................................59 Spiders and other related groups ..............................................60 Crustacea ................................................................62 Millipedes and centipedes ...................................................63 Hexapods (Hexapoda) ......................................................64 Bats (Chiroptera) ..........................................................67 Other vertebrates ..........................................................68 Future of underground research . 69 References . 70 Acknowledgements With regard to the fact that this material is mainly based on the book of POKORNÝ and HOLEC (2009), we would like to thank here all who already participated on the creation of the cited book. Here we would like to thank especially to RNDr. Vlado Papáč for providing photos of springtails. We also thank to language agency Skřivánek s.r.o. for the English translantion of the main parts of the text and to Ing. Lenka Zoubková for the final language improvement. We also thank to both opponents for their valuable comments on manuscript. We would also like to thank our workplace, which allows us to explore this issue, especially within teaching activities. We also 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 later approved by University Development Fund (UDF, FRVŠ in Czech) project No. 305/2010/F4/a – grant of the Ministry of Education, Youth and Sports (MEYS, MŠMT in Czech) and project of the Internal Foundation of UJEP – ”Paleontological research of the pseudokarst in north Bohemia”. Subterranean Habitats Introduction The authors have been paying considerable attention to the caves in the North Bohemian neovolca- nites for several years. The issue includes not only mapping caves in the field, but also the acquisition of other more detailed characteristics (part of data was published in POKORNÝ and HOLEC 2009). The colonization of caves by invertebrates was studied, althought data have not been still published. In the region, however, data on the occurrence of vertebrates, esp. bats, are available as well, e.g. BENDA and CHVÁTAL (2011). This was the background for the idea to create a compulsory school course, especially for foreign students of Jan Evangelista Purkyně University – Faculty of the Environment in Ústí nad Labem, which would call attention to the general scientific value of less known subterrannean environment. In this text we generalised our former unpublished text for students oriented mainly on regional lo- calities. The presented material brings the basic general information about the terrestrial caves and other natural terrestrial subterranean habitats. Water habitats, including interstitial habitats, were not mentioned because this text will serve as theoretical background for terrain excursion with students of Faculty of the Environment, Jan Evangelista Purkyně University in Ústí nad Labem. Our field trips with students will be oriented to the localities with terrestrial caves only and description of wa- ter habitats and water biota would stay without practical usage. For the same reason we prefer, for example, description of epigenetic pseudokarst. Description of other cave types, rarely surrounded our workplace or dangerous for excursions with students, was neglected. 6 Emergence of Selected Underground Spaces Emergence of Selected Underground Spaces Processes leading to the emergence of spaces below the surface of the earth can be divided into two basic groups – chemical and physical (= mechanical). By chemical processes we mean those which change the essence of the chemical substance of the primary rock. Generally these processes involve dissolution or oxidation. In the case of dissolution, the rock is dissolved by the action of a suitable sol- vent, such as water or a weak naturally occurring acid. This creates underground cavities of varying size and layout. As the dissolution equation generally works both ways, the reverse can happen, with dissolved substances accreting to create typical cave formations, known as speleothems. The most common chemical process leading to the creation of underground spaces is karstification, the result of which is karst, a characteristic type of landscape typified by the presence of surface and underground phenomena. The condition for this is the reaction of rainwater and other CO2-enriched surface waters with the rock (generally carbonate in composition – limestone, marble, dolomites) on the surface of the earth. Water travels down deep through fissures, where it dissolves the rock to form chimneys, passages and caves. In contrast to established opinion, karst phenomena do not only occur in limestone, but may be present in all rocks that have been at least partially dissolved. In the case of carbonates, the basic matter is dissolved according to the following schema (equations I, II) (PŘIBYL et al. 1992). + + 2– (I) CO2 + H2O ↔ H + (HCO3) – ↔ H + CO3 - 2+ 2- (II) (HCO3) + CaCO3 ↔ Ca + 2(HCO3) The work of CIGNA (1978) states that when describing the creation of the natural underground, we must base our arguments on the so-called three-component equilibrium: - CaCO3 (calcium carbonate, calcite) – the basic building block of calcareous rocks; in the broader sense any dissolved parent rock is considered to be a reaction component - water – a medium with mechanical and chemical action - - CO2 (carbon dioxide) – in the form of ions (HCO3) in a water solution acts as a kind of solvent CIGNA divides up the different types of formation depending on the number of components in- volved in the creation of the cave. 7 Subterranean Habitats Tab. I. Division of karst and non-karst