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06-Life at Extremes1 16 Caves and Karst Environments Natuschka M. Lee,1 Daniela B. Meisinger,1 Roman Aubrecht,2 Lubomir Kovacik,3 Cesareo Saiz-Jimenez,4 Sushmitha Baskar,5 Ramanathan Baskar,5 Wolfgang Liebl,1 Megan L. Porter6 and Annette Summers Engel7 1Department of Microbiology, Technische Universität München, Freising, Germany; 2Department of Geology and Palaeontology, Comenius University, Bratislava, Slovakia; 3Department of Botany, Comenius University, Bratislava, Slovakia; 4Instituto de Recursos Naturales y Agrobiologia, IRNAS-CSIC, Sevilla, Spain; 5Department of Environmental Science and Engineering, Guru Jambheshwar University of Science and Technology, Hisar, India; 6Department of Biological Sciences, University of Maryland, Baltimore, USA; 7Department of Geology and Geophysics, Louisiana State University, Baton Rouge, USA; now at: Department of Earth and Planetary Sciences, University of Tennessee, Knoxville, USA 16.1 Introduction a variety of different types of caves and intriguing cave creatures have been discov- Caves have played a fascinating role through- ered. Therefore, cave-based sciences play out the history of our planet and of our cul- an important role in enhancing our under- ture in different ways. Our first associations standing of the history of our planet and with caves centre on how they served as also form a foundation for exploring novel primitive dwelling sites for animals and concepts about the boundaries of life and the humans, and as settings for mysterious fan- evolution of extreme dark life ecosystems on tasies and myths (Fig. 16.1). Today, scien- Earth, as well as in other parts of the Universe tific exploration has added another aspect (Krajick, 2001; NOVA, 2002; SPACE/Malik to our connection to caves by revealing a and Writer, 2005; Forti, 2009). plethora of unexpected insights into diverse disciplines, including the natural sciences (geology, palaeontology, climatology, phys- 16.2 Description of Caves: Definition, ics, chemistry, biology and cosmology), medical sciences and engineering, as well Distribution and Biogeochemistry as social disciplines such as archaeology, theology, and the cultural history of man- 16.2.1 Introduction kind (Fig. 16.2). The foundation for this is the extreme nature of caves, characterized Many different definitions have been used by a lack of light and geographic isolation, to describe a cave or a cavern. The most but also by nutrient limitation and a range general way to describe a cave, irrespective of extreme redox conditions. In recent years, of its geological history and location, is to © CAB International 2012. Life at Extremes: Environments, 320 Organisms and Strategies for Survival (ed. E.M. Bell) Caves and Karst Environments 321 Fig. 16.1. Entrance zone of a cave: Cascade Caverns, Carter Caves State Resort Park, Kentucky, USA. Depending on the entrance morphology, sunlight only penetrates a limited distance into the cave. © Annette S. Engel. simply define it as a natural cavity in a rocky hundreds of kilometres in length. Only a environment where at least some part of it is few of these are accessible to humans. In fact, in total darkness. The science of exploration many cave openings consist only of micro- of caves and various karst features is termed scopic fractures. Despite numerous caving speleology (Gunn, 2004). Speleology is a activities all over the world, it is estimated broad interdisciplinary science; a multitude that, even in well explored areas like Europe of parameters and disciplines needs to be and North America, so far only 50% of all considered for a thorough exploration of caves in these regions have been accessed; the development of caves and their various globally, only ~10% of all caves have been impacts on their surroundings worldwide discovered (Eavis, 2009; Engel, 2011). The throughout history. subsurface can be regarded as one of the least Caves are developed in soluble rocks explored environment types on Earth, second and constitute a characteristic feature of only to the deep oceans. Caves often serve as karst (carbonate rock, such as limestone and the only available natural entrances and con- dolomite) and pseudokarst (non-carbonate nections to the subsurface, offering fascinating rock) landscapes that covers roughly windows into this vast and unexplored habitat. 15–20% of the Earth’s ice-free land surface Because of the increasing interest in cave (Ford and Williams, 2007). Although caves sciences and all the promising prospects they are found in various regions of the Earth and offer, plus the development of improved cav- at all latitudes, caves are generally not inter- ing technology that allows access to even the connected over large physiographic prov- most difficult caves, many novel types of inces, limited by the extent of rock type. caves have been discovered during the last Caves come in a wide range of shapes and decade (Eavis, 2009). Son Doong Cave, sizes, from micro-fissures to caverns several Vietnam, found in 2009, contains the largest thousands of metres deep and high, and discovered cavern to date, measuring up to 322 N.M. Lee et al. (a) (b) Fig. 16.2. (a) Grotta dei Cervi, Italy (discovered in February 1970), with black Neolithic wall paintings, made with bat guano. © Cesareo Saiz-Jimenez. (b) Castle in front of cave, Predjamski Grad, Slovenia. © Annette S. Engel. 140 by 140 m and 4.5 km in length. The cave known, and was discovered in Peru deepest cave, Krubera Cave, Georgia, near in 2004 at an altitude of nearly 5000 m. the Black Sea, was discovered in 2001 and Several fascinating underwater caves have extends to a depth of −2191 m into the sub- also been described, such as those in the surface. Qaqa Mach’ay Cave is the highest Yucatan Peninsula, Mexico, and the Nullarbor Caves and Karst Environments 323 Plain, Australia; each area contains hun- However, it is not only the size and dreds of kilometres of submerged passage- depth of caves that are impressive (Figs 16.3 ways. However, even caves that have been and 16.4); many of their contents undisput- known about and explored for many years, edly continue to capture our imagination e.g. Mammoth Cave, Kentucky, USA, also and curiosity, e.g. their spectacular spele- continue to reveal astonishing insights. othems (cave rock and mineral formations), Due to natural karstification as well as such as stalactites, stalagmites and giant continued exploration, Mammoth Cave is crystals that may be over 70 m in length, now estimated to cover a distance of around and fascinating creatures belonging to 600 km, and as such represents the longest unique, extreme ecosystems (Taylor, 1999; cave known on Earth (for a summary of Krajick, 2001; Culver and Pipan, 2009; Eavis, spectacular cave types, see NSS GEO2 com- 2009, http://www.canyonsworldwide.com/ mittee, 2011). crystals/mainframe3.html). (a) (b) (c) (d) Fig. 16.3. Examples of spectacular constructions in different types of caves. (a) The large room of Škocjan Caves Regional Park, Slovenia, a UNESCO site. This photo highlights the massive size that speleothems can reach (people for scale in the lower left). © Annette S. Engel. (b) Unique formations of unknown nature (possibly biospeleothems) inside a sandstone cave in Venezuela (Aubrecht et al., 2008). © Jan Schloegl and Roman Aubrecht. (c) Entrances to caves can be very small and embedded in water; Cascade Caverns, Carter Caves State Resort Park, Kentucky, USA. © Annette S. Engel. (d) Example of different colour formations due to various microbiological redox processes. Sulfidic spring in a cave with toxic hydrogen sulfidic gases, Lower Kane Cave, Wyoming, USA. © Annette S. Engel. 324 N.M. Lee et al. (a) (b) (c) (d) Fig. 16.4. Different types of speleothems and biospeleothems in a sandstone cave in Venezuela. (a–c) sandstone cave in Venezuela, © Roman Aubrecht. (d) Driny Cave in Slovakia, © Lubomir Kovacik. 16.2.2 Classification and formation of caves Although new caves are constantly being formed somewhere on Earth, the Caves are classified using several criteria majority of the caves so far described and (Northup and Lavoie, 2001; Gunn, 2004; explored have most likely persisted for Engel, 2011): the solid rock/bedrock type, longer periods, from thousands to millions the proximity to the groundwater table, the of years. Irrespective of their speleogenetic overall passage morphology and organiza- histories, all caves are constantly changing tion (i.e. size, length, depth, routes of fluid over time, either by natural karstification flow, etc.) and the speleogenetic history processes or simply due to continued explo- (related to the origin and development of ration. Depending on the proximity to the caves). So far, at least 250 different minerals groundwater table, two different general have been described from karst and pseu- modes of cave formation can be discerned: dokarst settings (Hill and Forti, 1997). (i) epigenic caves, as the most commonly However, the most common rock types are described caves, are related to surface pro- calcareous rocks (e.g. limestone, which cesses by formation at or proximal to the underlies about 15% of Earth’s surface) and groundwater table, and are critically depend- basaltic rock (e.g. lava tubes). Other less ent on the hydrological conditions of the common rock types include other volcanic region; and (ii) hypogenic caves, which are deposits, gypsum, granite, quartzite, sand- related to subsurface processes because they stone, salt and even ice. form by the action of rising fluids, such as Caves and Karst Environments 325 water or gases, at or below the water table 3. Volcanism leading to lava flows that (Engel, 2011). The most commonly described produce lava tube caves (Halliday, 2004), hypogenic caves to date are sulfidic caves, which first may result in sterilized surfaces which are formed by a combination of abi- after a volcanic eruption that may then be otically and microbially produced sulfuric colonized through time. Examples of exten- acid (Engel, 2007). Irrespective of the cave’s sive lava tube caves have been described for relation to surface or subsurface processes, the Kilauea Volcano, Hawaii, USA.
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