Application of the gravitydriven regional groundwater flow (GDRGF) concept to the hydrogeologically complex thick carbonate system of the Transdanubian Range (TR), Hungary, is justified based on the principle of hydraulic continuity. The GDRGF concept informs about basin hydraulics and groundwater as a geologic agent. It became obvious that the effect of heterogeneity and anisotropy on the flow pattern could be derived from hydraulic reactions of the aquifer system. The topography and heat as driving forces were examined by numerical simulations of flow and heat transport. Evaluation of groups of springs, in terms of related discharge phenomena and regional chloride distribution, reveals the dominance of topographydriven flow when considering flow and related chemical and temperature patterns. Moreover, heat accumulation beneath the confined part of the system also influences these patterns. The presence of cold, lukewarm and thermal springs and related wetlands, creeks, mineral precipitates, and epigenic and hypogenic caves validates the existence of GDRGF in the system. Vice versa, groups of springs reflect rock–water interaction and advective heat transport and inform about basin hydraulics. Based on these findings, a generalized conceptual GDRGF model is proposed for an unconfined and confined carbonate region. An interface was revealed close to the margin of the unconfined and confined carbonates, determined by the GDRGF and freshwater and basinal fluids involved. The application of this model provides a background to interpret manifestations of flowing groundwater in thick carbonates generally, including porosity enlargement and hydrocarbon and heat accumulation.

Homo species were exposed to a new biogeochemical environment when they began to occupy caves. Here we report the first evidence of palaeopollution through geochemical analyses of heavy metals in four renowned archaeological caves of the Iberian Peninsula spanning the last million years of human evolution. Heavy metal contents reached high values due to natural (guano deposition) and anthropogenic factors (e.g. combustion) in restricted cave environments. The earliest anthropogenic pollution evidence is related to Neanderthal hearths from Gorham's Cave (Gibraltar), being one of the first milestones in the so-called “Anthropocene”. According to its heavy metal concentration, these sediments meet the present-day standards of “contaminated soil”. Together with the former, the Gibraltar Vanguard Cave, shows Zn and Cu pollution ubiquitous across highly anthropic levels pointing to these elements as potential proxies for human activities. Pb concentrations in Magdalenian and Bronze age levels at El Pirulejo site can be similarly interpreted. Despite these high pollution levels, the contaminated soils might not have posed a major threat to Homo populations. Altogether, the data presented here indicate a long-term exposure of Homo to these elements, via fires, fumes and their ashes, which could have played certain role in environmental-pollution tolerance, a hitherto neglected influence.

Caves formed by rising sulfuric waters have been described from all over the world in a wide variety of climate  settings, from arid regions to mid-latitude and alpine areas. H2S is generally formed at depth by reduction of  sulfates in the presence of hydrocarbons and is transported in solution through the deep aquifers. In tectonically  disturbed areas major fractures eventually allow these H2S-bearing fluids to rise to the surface where oxidation  processes can become active producing sulfuric acid. This extremely strong acid reacts with the carbonate  bedrock creating caves, some of which are among the largest and most spectacular in the world. Production of  sulfuric acid mostly occurs at or close to the water table but also in subaerial conditions in moisture films and  droplets in the cave environment. These caves are generated at or immediately above the water table, where  condensation–corrosion processes are dominant, creating a set of characteristic meso- and micromorphologies.  Due to their close connection to the base level, these caves can also precisely record past hydrological and  geomorphological settings. Certain authigenic cave minerals, produced during the sulfuric acid speleogenesis  (SAS) phase, allow determination of the exact timing of speleogenesis. This paper deals with the morphological,  geochemical and mineralogical description of four very typical sulfuric acid water table caves in Europe: the  Grotte du Chat in the southern French Alps, the Acqua Fitusa Cave in Sicily (Italy), and the Bad Deutsch Altenburg  and Kraushöhle caves in Austria

Application of the gravity-driven regional  groundwater flow (GDRGF) concept to the  hydrogeologically complex thick carbonate system of the  Transdanubian Range (TR), Hungary, is justified based on  the principle of hydraulic continuity. The GDRGF concept  informs about basin hydraulics and groundwater as a  geologic agent. It became obvious that the effect of  heterogeneity and anisotropy on the flow pattern could be  derived from hydraulic reactions of the aquifer system.  The topography and heat as driving forces were examined  by numerical simulations of flow and heat transport.  Evaluation of groups of springs, in terms of related  discharge phenomena and regional chloride distribution,  reveals the dominance of topography-driven flow when  considering flow and related chemical and temperature  patterns. Moreover, heat accumulation beneath the confined  part of the system also influences these patterns. The  presence of cold, lukewarm and thermal springs and  related wetlands, creeks, mineral precipitates, and epigenic  and hypogenic caves validates the existence of GDRGF in  the system. Vice versa, groups of springs reflect rock–  water interaction and advective heat transport and inform  about basin hydraulics. Based on these findings, a  generalized conceptual GDRGF model is proposed for  an unconfined and confined carbonate region. An interface  was revealed close to the margin of the unconfined and  confined carbonates, determined by the GDRGF and  freshwater and basinal fluids involved. The application  of this model provides a background to interpret manifestations  of flowing groundwater in thick carbonates  generally, including porosity enlargement and hydrocarbon  and heat accumulation.

Dissolution of fractured rocks is often accompanied by the formation of highly localized flow paths. While the fluid flow follows existing fractures in the rock, these fissures do not, in general, open uniformly. Simulations and laboratory experiments have shown that distinct channels or “wormholes”develop within the fracture, from which a single highly localized flow path eventually emerges. The aim of the present work is to investigate how these emerging flow paths are influenced by the initial aperture field. We have simulated the dissolution of a single fracture starting from a spatially correlated aperture distribution. Our results indicate a surprising insensitivity of the evolving dissolution patterns and flow rates to the amplitude and correlation length characterizing the imposed aperture field. We connect the similarity in outcomes to the self-organization of the flow into a small number of wormholes, with the spacing determined of the longest wormholes. We have also investigated the effect of a localized region of increased aperture on the developing dissolution patterns. A competition was observed between the tendency of the high-permeability region to develop the dominant wormhole and the tendency of wormholes to spontaneously nucleate throughout the rest of the fracture. We consider the consequences of these results for the modeling of dissolution in fractured and porous rocks.

Karst on the Nullarbor Plain has been studied and described in detail in the past, but it lacked the determination of the karst discharge and palaeo-watertable levels that would explain the palaeohydrological regime in this area. This study explores the existence of previously unrecognised features in this area – karst pocket valleys – and gives a review on pocket valleys worldwide. Initial GIS analyses were followed up by detailed field work, sampling, mapping and measuring of morphological, geological, and hydrological characteristics of representative
valleys on the Wylie and Hampton scarps of the Nullarbor Plain. Rock and sand samples were examined for mineralogy, texture and grain size, and a U–Pb dating of a speleothem froma cave within a pocket valley enabled the establishment of a time frame of the pocket valleys formation and its palaeoenvironmental implications. The pocket valleys document the hydrological evolution of the Nullarbor karst system and the Neogene–Pleistocene palaeoclimatic evolution of the southern hemisphere. A review of pocket valleys in different climatic and geological settings suggests that their basic characteristics remain the same, and their often overlooked utility as environmental indicators can be used for further palaeoenvironmental studies. The main period of intensive karstification and widening of hydrologically active underground conduits is placed into the wetter climates of the Pliocene epoch. Subsequent drier climates and lowering of the watertable that followed sea-level retreat in the Quaternary resulted in formation of the pocket valleys (gravitational undermining, slumping, exudation and collapse), which, combined with periodic heavy rainfall events and discharge due to impeded drainage, caused the retreat of the pocket valleys from the edge of escarpments.

Speleogenesis is the development of well-organized cave systems by fluids moving through fissures of a soluble rock. Epigenic caves induced by biogenic CO2 soil production are dominant, whereas hypogenic caves resulting from uprising deep flow not directly connected to adjacent recharge areas appear to be more frequent than previously considered. The conceptual models of epigenic cave development moved from early models, through the “four-states model” involving fracture influence to explain deep loops, to the digital models demonstrating the adjustment of the main flow to the water table. The relationships with base level are complex and cave levels must be determined from the elevation of the vadose-phreatic transitions. Since flooding in the epiphreatic zone may be important, the top of the loops in the epiphreatic zone can be found significantly high above the base level. The term Paragenesis is used to describe the upward development of conduits as their lower parts fill with sediments. This process often records a general baselevel rise. Sediment influx is responsible for the regulation of long profiles by paragenesis and contributes to the evolution of profiles from looping to water table caves. Dating methods allow identification of the timing of cave level evolution. The term Ghost-rock karstification is used to describe a 2-phase process of speleogenesis, with a first phase of partial solution of rock along fractures in low gradient conditions leaving a porous matrix, the ghost-rock, then a second phase of mechanical removing of the ghost-rock mainly by turbulent flow in high gradient conditions opening the passages and forming maze caves. The first weathering phase can be related either to epigenic infiltration or to hypogenic upflow, especially in marginal areas of sedimentary basins. The vertical pattern of epigenic caves is mainly controlled by timing, geological structure, types of flow and base-level changes. We define several cave types as (1) juvenile, where they are perched above underlying aquicludes; (2) looping, where recharge varies greatly with time, to produce epiphreatic loops; (3) water-table caves where flow is regulated by a semi-pervious cover; and (4) caves in the equilibrium stage where flow is transmitted without significant flooding. Successive base-level drops caused by valley entrenchment make cave levels, whereas baselevel rise is defined in the frame of the Per ascensum Model of Speleogenesis (PAMS), where deep passages are flooded and drain through vauclusian springs. The PAMS can be active after any type of baselevel rise (transgression, fluvial aggradation, tectonic subsidence) and explains most of the deep phreatic cave systems except for hypogenic.

The term Hypogenic speleogenesis is used to describe cave development by deep upflow independent of adjacent recharge areas. Due to its deep origin, water frequently has a high CO2-H2S concentration and a thermal anomaly, but not systemati­cally. Numerous dissolution processes can be involved in hypogenic speleogenesis, which often include deep-seated acidic sources of CO2 and H2S, “hydrothermal” cooling, mixing corrosion, Sulfuric Acid Speleogenesis (SAS), etc. SAS particularly involves the condensation-corrosion processes, resulting in the fast expansion of caves above the water table, i.e. in an atmo­spheric environment. The hydrogeological setting of hypogenic speleogenesis is based on the Regional Gravity Flow concept, which shows at the basin scales the sites of convergences and upflows where dissolution focuses. Each part of a basin (mar­ginal, internal, deep zone) has specific conditions. The coastal basin is a sub-type. In deformed strata, flow is more complex according to the geological structure. However, upflow and hypogenic speleogenesis concentrate in structural highs (buried anticlines) and zones of major disruption (faults, overthrusts). In disrupted basins, the geothermal gradient “pumps” the me­teoric water at depth, making loops of different depths and characteristics. Volcanism and magmatism also produce deep hypogenic loops with “hyperkarst” characteristics due to a combination of deep-seated CO2, H2S, thermalism, and microbial activity. In phreatic conditions, the resulting cave patterns

can include geodes, 2–3D caves, and giant ascending shafts. Along the water table, SAS with thermal air convection induces powerful condensation-corrosion and the development of upwardly dendritic caves, isolated chambers, water table sulfuricacid caves. In the vadose zone, “smoking” shafts evolve under the influence of geothermal gradients producing air convectionand condensation-corrosion.

Likely future directions for research will probably involve analytical and modeling methods, especially using isotopes, dating, chemical simulations, and field investigations focused on the relationships between processes and resulting morphologies.

Speleogenesis is the development of well-organized cave systems by fluids moving through fissures of a soluble rock. Epigenic caves induced by biogenic CO2 soil production are dominant, whereas hypogenic caves resulting from uprising deep flow not directly connected to adjacent recharge areas appear to be more frequent than previously considered. The conceptual models of epigenic cave development moved from early models, through the “four-states model” involving fracture influence to explain deep loops, to the digital models demonstrating the adjustment of the main flow to the water table. The relationships with base level are complex and cave levels must be determined from the elevation of the vadose-phreatic transitions. Since flooding in the epiphreatic zone may be important, the top of the loops in the epiphreatic zone can be found significantly high above the base level. The term Paragenesis is used to describe the upward development of conduits as their lower parts fill with sediments. This process often records a general baselevel rise. Sediment influx is responsible for the regulation of long profiles by paragenesis and contributes to the evolution of profiles from looping to water table caves. Dating methods allow identification of the timing of cave level evolution. The term Ghost-rock karstification is used to describe a 2-phase process of speleogenesis, with a first phase of partial solution of rock along fractures in low gradient conditions leaving a porous matrix, the ghost-rock, then a second phase of mechanical removing of the ghost-rock mainly by turbulent flow in high gradient conditions opening the passages and forming maze caves. The first weathering phase can be related either to epigenic infiltration or to hypogenic upflow, especially in marginal areas of sedimentary basins. The vertical pattern of epigenic caves is mainly controlled by timing, geological structure, types of flow and base-level changes. We define several cave types as (1) juvenile, where they are perched above underlying aquicludes; (2) looping, where recharge varies greatly with time, to produce epiphreatic loops; (3) water-table caves where flow is regulated by a semi-pervious cover; and (4) caves in the equilibrium stage where flow is transmitted without significant flooding. Successive base-level drops caused by valley entrenchment make cave levels, whereas baselevel rise is defined in the frame of the Per ascensum Model of Speleogenesis (PAMS), where deep passages are flooded and drain through vauclusian springs. The PAMS can be active after any type of baselevel rise (transgression, fluvial aggradation, tectonic subsidence) and explains most of the deep phreatic cave systems except for hypogenic.

The term Hypogenic speleogenesis is used to describe cave development by deep upflow independent of adjacent recharge areas. Due to its deep origin, water frequently has a high CO2-H2S concentration and a thermal anomaly, but not systemati­cally. Numerous dissolution processes can be involved in hypogenic speleogenesis, which often include deep-seated acidic sources of CO2 and H2S, “hydrothermal” cooling, mixing corrosion, Sulfuric Acid Speleogenesis (SAS), etc. SAS particularly involves the condensation-corrosion processes, resulting in the fast expansion of caves above the water table, i.e. in an atmo­spheric environment. The hydrogeological setting of hypogenic speleogenesis is based on the Regional Gravity Flow concept, which shows at the basin scales the sites of convergences and upflows where dissolution focuses. Each part of a basin (mar­ginal, internal, deep zone) has specific conditions. The coastal basin is a sub-type. In deformed strata, flow is more complex according to the geological structure. However, upflow and hypogenic speleogenesis concentrate in structural highs (buried anticlines) and zones of major disruption (faults, overthrusts). In disrupted basins, the geothermal gradient “pumps” the me­teoric water at depth, making loops of different depths and characteristics. Volcanism and magmatism also produce deep hypogenic loops with “hyperkarst” characteristics due to a combination of deep-seated CO2, H2S, thermalism, and microbial activity. In phreatic conditions, the resulting cave patterns

can include geodes, 2–3D caves, and giant ascending shafts. Along the water table, SAS with thermal air convection induces powerful condensation-corrosion and the development of upwardly dendritic caves, isolated chambers, water table sulfuricacid caves. In the vadose zone, “smoking” shafts evolve under the influence of geothermal gradients producing air convectionand condensation-corrosion.

Likely future directions for research will probably involve analytical and modeling methods, especially using isotopes, dating, chemical simulations, and field investigations focused on the relationships between processes and resulting morphologies.

Speleogenesis is the development of well-organized cave systems by fluids moving through fissures of a soluble rock. Epigenic caves induced by biogenic CO2 soil production are dominant, whereas hypogenic caves resulting from uprising deep flow not directly connected to adjacent recharge areas appear to be more frequent than previously considered. The conceptual models of epigenic cave development moved from early models, through the “four-states model” involving fracture influence to explain deep loops, to the digital models demonstrating the adjustment of the main flow to the water table. The relationships with base level are complex and cave levels must be determined from the elevation of the vadose-phreatic transitions. Since flooding in the epiphreatic zone may be important, the top of the loops in the epiphreatic zone can be found significantly high above the base level. The term Paragenesis is used to describe the upward development of conduits as their lower parts fill with sediments. This process often records a general baselevel rise. Sediment influx is responsible for the regulation of long profiles by paragenesis and contributes to the evolution of profiles from looping to water table caves. Dating methods allow identification of the timing of cave level evolution. The term Ghost-rock karstification is used to describe a 2-phase process of speleogenesis, with a first phase of partial solution of rock along fractures in low gradient conditions leaving a porous matrix, the ghost-rock, then a second phase of mechanical removing of the ghost-rock mainly by turbulent flow in high gradient conditions opening the passages and forming maze caves. The first weathering phase can be related either to epigenic infiltration or to hypogenic upflow, especially in marginal areas of sedimentary basins. The vertical pattern of epigenic caves is mainly controlled by timing, geological structure, types of flow and base-level changes. We define several cave types as (1) juvenile, where they are perched above underlying aquicludes; (2) looping, where recharge varies greatly with time, to produce epiphreatic loops; (3) water-table caves where flow is regulated by a semi-pervious cover; and (4) caves in the equilibrium stage where flow is transmitted without significant flooding. Successive base-level drops caused by valley entrenchment make cave levels, whereas baselevel rise is defined in the frame of the Per ascensum Model of Speleogenesis (PAMS), where deep passages are flooded and drain through vauclusian springs. The PAMS can be active after any type of baselevel rise (transgression, fluvial aggradation, tectonic subsidence) and explains most of the deep phreatic cave systems except for hypogenic.

The term Hypogenic speleogenesis is used to describe cave development by deep upflow independent of adjacent recharge areas. Due to its deep origin, water frequently has a high CO2-H2S concentration and a thermal anomaly, but not systemati­cally. Numerous dissolution processes can be involved in hypogenic speleogenesis, which often include deep-seated acidic sources of CO2 and H2S, “hydrothermal” cooling, mixing corrosion, Sulfuric Acid Speleogenesis (SAS), etc. SAS particularly involves the condensation-corrosion processes, resulting in the fast expansion of caves above the water table, i.e. in an atmo­spheric environment. The hydrogeological setting of hypogenic speleogenesis is based on the Regional Gravity Flow concept, which shows at the basin scales the sites of convergences and upflows where dissolution focuses. Each part of a basin (mar­ginal, internal, deep zone) has specific conditions. The coastal basin is a sub-type. In deformed strata, flow is more complex according to the geological structure. However, upflow and hypogenic speleogenesis concentrate in structural highs (buried anticlines) and zones of major disruption (faults, overthrusts). In disrupted basins, the geothermal gradient “pumps” the me­teoric water at depth, making loops of different depths and characteristics. Volcanism and magmatism also produce deep hypogenic loops with “hyperkarst” characteristics due to a combination of deep-seated CO2, H2S, thermalism, and microbial activity. In phreatic conditions, the resulting cave patterns

can include geodes, 2–3D caves, and giant ascending shafts. Along the water table, SAS with thermal air convection induces powerful condensation-corrosion and the development of upwardly dendritic caves, isolated chambers, water table sulfuricacid caves. In the vadose zone, “smoking” shafts evolve under the influence of geothermal gradients producing air convectionand condensation-corrosion.

Likely future directions for research will probably involve analytical and modeling methods, especially using isotopes, dating, chemical simulations, and field investigations focused on the relationships between processes and resulting morphologies.

Speleogenesis is the development of well-organized cave systems by fluids moving through fissures of a soluble rock. Epigenic caves induced by biogenic CO2 soil production are dominant, whereas hypogenic caves resulting from uprising deep flow not directly connected to adjacent recharge areas appear to be more frequent than previously considered. The conceptual models of epigenic cave development moved from early models, through the “four-states model” involving fracture influence to explain deep loops, to the digital models demonstrating the adjustment of the main flow to the water table. The relationships with base level are complex and cave levels must be determined from the elevation of the vadose-phreatic transitions. Since flooding in the epiphreatic zone may be important, the top of the loops in the epiphreatic zone can be found significantly high above the base level. The term Paragenesis is used to describe the upward development of conduits as their lower parts fill with sediments. This process often records a general baselevel rise. Sediment influx is responsible for the regulation of long profiles by paragenesis and contributes to the evolution of profiles from looping to water table caves. Dating methods allow identification of the timing of cave level evolution. The term Ghost-rock karstification is used to describe a 2-phase process of speleogenesis, with a first phase of partial solution of rock along fractures in low gradient conditions leaving a porous matrix, the ghost-rock, then a second phase of mechanical removing of the ghost-rock mainly by turbulent flow in high gradient conditions opening the passages and forming maze caves. The first weathering phase can be related either to epigenic infiltration or to hypogenic upflow, especially in marginal areas of sedimentary basins. The vertical pattern of epigenic caves is mainly controlled by timing, geological structure, types of flow and base-level changes. We define several cave types as (1) juvenile, where they are perched above underlying aquicludes; (2) looping, where recharge varies greatly with time, to produce epiphreatic loops; (3) water-table caves where flow is regulated by a semi-pervious cover; and (4) caves in the equilibrium stage where flow is transmitted without significant flooding. Successive base-level drops caused by valley entrenchment make cave levels, whereas baselevel rise is defined in the frame of the Per ascensum Model of Speleogenesis (PAMS), where deep passages are flooded and drain through vauclusian springs. The PAMS can be active after any type of baselevel rise (transgression, fluvial aggradation, tectonic subsidence) and explains most of the deep phreatic cave systems except for hypogenic.

The term Hypogenic speleogenesis is used to describe cave development by deep upflow independent of adjacent recharge areas. Due to its deep origin, water frequently has a high CO2-H2S concentration and a thermal anomaly, but not systemati­cally. Numerous dissolution processes can be involved in hypogenic speleogenesis, which often include deep-seated acidic sources of CO2 and H2S, “hydrothermal” cooling, mixing corrosion, Sulfuric Acid Speleogenesis (SAS), etc. SAS particularly involves the condensation-corrosion processes, resulting in the fast expansion of caves above the water table, i.e. in an atmo­spheric environment. The hydrogeological setting of hypogenic speleogenesis is based on the Regional Gravity Flow concept, which shows at the basin scales the sites of convergences and upflows where dissolution focuses. Each part of a basin (mar­ginal, internal, deep zone) has specific conditions. The coastal basin is a sub-type. In deformed strata, flow is more complex according to the geological structure. However, upflow and hypogenic speleogenesis concentrate in structural highs (buried anticlines) and zones of major disruption (faults, overthrusts). In disrupted basins, the geothermal gradient “pumps” the me­teoric water at depth, making loops of different depths and characteristics. Volcanism and magmatism also produce deep hypogenic loops with “hyperkarst” characteristics due to a combination of deep-seated CO2, H2S, thermalism, and microbial activity. In phreatic conditions, the resulting cave patterns

can include geodes, 2–3D caves, and giant ascending shafts. Along the water table, SAS with thermal air convection induces powerful condensation-corrosion and the development of upwardly dendritic caves, isolated chambers, water table sulfuricacid caves. In the vadose zone, “smoking” shafts evolve under the influence of geothermal gradients producing air convectionand condensation-corrosion.

Likely future directions for research will probably involve analytical and modeling methods, especially using isotopes, dating, chemical simulations, and field investigations focused on the relationships between processes and resulting morphologies.

The aim of this study is to characterize in detail, the mineralogy of different-shaped concretions as well as to investigate the physico-chemical parameters of the associated mine drainage and drip waters in the Santa Barbara level of the Libiola Mine (NW Italy) by several geochemical and mineralogical techniques. Under the term “minothems” we are grouping all those secondary minerals that occur under certain form or shape related to the conditions under which they formed but occur in a mine, or in any artificial underground environment (i.e., "mine speleothems"). Different types of minothems (soda straw stalactites, stalactites, and draperies) were sampled and analyzed. Mineralogical results showed that all the samples of stalactites, stalagmite and draperies are characterized by poorly crystalline goethite. There are significant differences either in their texture and chemistry. Stalactites are enriched in Zn, Cd, and Co in respect to other minothems and show botryoidal textures; some of these exhibit a concentric layering marked by the alternation of botryoidal and fibrous-radiating textures; the draperies are enriched in V and show aggregates of sub-spheroidal goethite forming compact mosaic textures. Geochemical investigations show that the composition and physico-chemical parameters of mine drainage and drip waters are different from the other acidic mine water occurrences in different areas of the Libiola Mine, where minothems are less abundant. All mine water samples contain Cu, Ni, and Zn in appreciable levels, and the physico-chemical conditions are consistent with the stability of ferrihydrite, which however tends to transform into goethite upon ageing.

Karst environments can be grouped into three broad categories, based on their vertical position in the landscape. There are surface habitats, ones exposed to light; there are shallow subterranean (aphotic) habitats oft en with small to intermediate sized spaces; there are deep subterranean habitats (caves) with large sized spaces. Faunal records are most complete for caves, and on a global basis, more than 10,000 species are limited to this habitat. Hundreds of other species, especially bats, depend on caves for some part of their life cycle. A large, but most unknown number of species are limited to shallow subterranean habitats in karst, such as epikarst and the milieu souterrain superficiel. Species in both these categories of habitats typically show a number of morphological adaptations for life in darkness, including loss of eyes and pigment, and elaboration of extra-optic sensory structures. Surface habitats, such as sinkholes, karst springs, thin soils, and rock faces, are habitats, but not always recognized as karst habitats. Both aphotic karst habitats and twilight habitats (such as open air pits) may serve as important temporary refuges for organisms avoiding temperature extremes on the surface.