Ajalon cave

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Ayalon Cave (Israel)
Ajalon cave
Ajalon cave
Beer Sheva
Beer Sheva
Jerusalem
Jerusalem
Carmel
Carmel
Tel Aviv-Jaffa
Tel Aviv-Jaffa
The location of the Ajalon Cave in Israel. From about Carmel according Beersheba runs the Yarkon Taninim Aquifer

The Ajalon Cave ( Hebrew מערת איילון) near Ramla , between Tel Aviv and Jerusalem , is the second largest limestone cave in Israel with a length of more than 2,700 meters . Her name is derived from her location in the valley of Ajalon , mentioned in the Old Testament ( Jos 10,12  EU ) . The Ajalon Cave was discovered on April 24, 2006 while working in the quarry of a cement factory. It should remain inaccessible to the public to protect the unique fauna with at least six endemic species of arthropods . The operator of the quarry stated that regardless of possible disruptions to operations, his company is interested in preserving the cave and its ecosystem.

Location and discovery

Northern edge of the Nesher quarry, cross-section with the Ajalon cave

The Ajalon Cave is located about four kilometers southeast of the city of Ramla and 21 kilometers from the Mediterranean Sea in the Israeli central district (coordinates: 31 ° 54 ′ 37.5 ″  N , 34 ° 55 ′ 39.3 ″  E ). It is located in a limestone quarry owned by Nesher Israel Cement Enterprises Ltd., which has an area of ​​around 1300 × 600 meters. The bottom of the quarry was about 100 meters below the original terrain level in 2006, and thus below the groundwater level of the 1950s. Routine investigations of the subsurface revealed indications of voids in the rock. Upon closer examination, members of the Hebrew University of Jerusalem and volunteers from the Israel Cave Research Center determined the extent of the cave system.

geology

Connecting passage in the upper level of the cave

Structure of the cave

Despite the small size of the country, Israel is extraordinarily rich in caves, most of which are karst caves . In the karst there is a high risk of sudden collapse of underground cavities, both in quarries and in settlements on insufficiently investigated soil. In the past, there were repeated break-ins in the area of ​​the Ayalon Saline Anomaly , even in the immediate vicinity of inhabited houses. In December 1997, a fatal accident occurred in the quarry of Nesher Israel Cement Enterprises. The ceiling of a cavity had become so thin as a result of the dismantling work that a bulldozer and the driver fell 40 meters into the depth. Since then, numerous drillings have been carried out in the quarry during the mining operation in order to identify danger spots at an early stage. In the process, cavities were repeatedly discovered on the northern and eastern edges of the quarry, in the vicinity of the Ajalon cave, but they were not as large and some had already collapsed.

The Ajalon Cave is a karst cave in the Upper Cretaceous Israel, which consists of a number of narrow, partly vertical passages with a total length of 2,780 meters. The cave extends on two main levels between 11.30 and 49 meters above sea ​​level and occupies an area of ​​only about 100 × 140 meters. The upper level is 40 to 49 meters above sea level. It consists primarily of narrow, labyrinthine interconnected corridors with a round or elliptical cross-section and a diameter of 0.30 to 1.40 meters. The total length of the tubes in the upper level is almost 2000 meters, which is about three quarters of the length of the cave. The tubes have partially collapsed or filled with clay . Part of the walls is covered with calcite crusts, which in some places have predominantly or completely closed the tubes.

The lower level is between 11.30 and 32 meters above sea level and is connected to the upper level by vertical shafts. It has wider passages and three chambers. The lower area is characterized by significantly more collapses than the upper level, making many passages impassable. The largest chamber is about 200 meters from the cave entrance and forms a hall more than 30 meters high with a diameter of 40 meters. The walls of the hall are covered with bright calcite crystals. On the lower part they form a layer about five centimeters thick, which becomes thinner towards the top.

There is a layer of rock that is only about 30 meters thick above the hall. Although the hall is free, calculations have shown that it is not very stable. This finding is supported by the fact that parts of the ceilings in the chambers have fallen down since the cavity was formed, and some corridors have been blocked by debris and deformations.

Part of the hall area was taken up by a cave lake that was around four meters deep in 2006 and contains salty groundwater with a high concentration of hydrogen sulfide . There are carts below the surface of the water , the formation of which is attributed to a descending water current. The analysis of the water showed a temperature of 28.5 to 30 ° C, 4.5 ppm hydrogen sulfide, a pH value of 6.8 and a salt content of 490 to 1300 mg / l of chloride in the lower areas  . The water of the lake is anoxic below one meter . The layer of sulphurous thermal water is covered by surface water with a temperature of only around 25 ° C, the properties of which largely correspond to the surrounding groundwater, and in which the crustaceans of the lake live.

Ajalon Cave, large chamber with the cave lake, in the foreground and in the back left debris from previous ceiling collapses

Hydrogeology

The Ajalon Cave is located in the area of ​​the Yarkon Taninim Aquifer , the most important freshwater reservoir in Israel. The Yarkon Taninim Aquifer takes its name from two springs fed by it. The Yarkon rises from freshwater springs about 15 kilometers east of Tel Aviv . The Taninim , on the other hand, draws salty water from springs around 60 kilometers to the north. The Yarkon Taninim aquifer is fed by precipitation in the mountainous regions of the West Bank and runs on the eastern edge of the Israeli coastal plain from Carmel in the north to south of Beersheba . Although it has been used and studied for almost a century, its exact structure remains unclear to this day. It is undisputed that it is formed from two isolated aquifers, each around 350 meters thick, which are usually considered together and thus form an aquifer 700 to 1000 meters thick.

The aquifer is threatened by intensive water abstraction. The water abstraction by Israel clearly exceeds the new formation of groundwater, so that since the 1950s there has been a drop in the groundwater level and, due to the smaller amount of water, an increase in the salt content in the groundwater. Another threat is the entry of nitrate from untreated sewage, both in Israel and the West Bank.

Sulfur and salty groundwater was discovered in the region as early as 1932; later, groundwater with an inexplicably high salinity was pumped from numerous wells in the area . The affected area, the Ayalon Saline Anomaly , covers an area of ​​200 square kilometers. Initially, geologists had assumed various possible causes on the earth's surface, including salt washed out from mineral-rich rock and residues of fertilizers.

The water conditions in the Nesher quarry and its surroundings were examined over a period of four years. By analyzing the water temperature, mineral content and concentration of hydrogen sulfide in the water from 68 boreholes in the region, it was established that the Ayalon Saline Anomaly , in the center of which the quarry with the Ajalon cave is located, is actually fed by thermal springs . In the closing sentence of their publication, which appeared a few weeks before the discovery of the Ajalon Cave, the researchers pointed out the dangers of large cavities under the earth's surface that should be taken into account during construction work and in quarries in the region.

Creation of the Ajalon cave

The Ajalon cave was created by the salt and sulfur-containing thermal water escaping in its area, which mixed with the existing groundwater and expanded existing cracks in the rock. Until a few decades ago, the lower level of the cave was flooded. The chemical composition of the water, the nature of the cave walls and the microfauna found lead to the conclusion that the process of cave formation is still in progress. In this respect, the Ajalon Cave is comparable to the Frasassi Caves in Italy and the Movile Cave in Romania. Such caves arise when the thermal water escaping hydrogen sulfide reacts with dissolved oxygen in the surrounding groundwater, or is oxidized to sulfuric acid by microbes . In the simplest case, the resulting sulfuric acid reacts with the surrounding limestone and converts it into gypsum and carbon dioxide :

In addition, other chemical reactions to varying degrees occur within the framework of bacterial metabolism and depending on the available substances. Sulfur, oxygen, carbon and nitrogen are regularly involved; some of these reactions also produce acids that attack the limestone.

Speleobiology

Catching of crustaceans in the lake of the Ajalon Cave

Environmental conditions

Until it opened, the Ajalon Cave formed an ecosystem that was closed off from the outside world , into which neither water nor organic material from the surface could penetrate because of a layer of limestone above it that was tens of meters thick. The air temperature in most areas of the cave fluctuates between 26 and 28 ° C with a humidity of more than 94%. The atmosphere in the lower level of the cave is high in hydrogen sulfide.

The organisms that existed in the cave were constantly dependent on the biomass produced by chemoautotrophic bacteria. Such ecosystems are rare worldwide, in Israel such a system was first described in 1968 with the source En Nur in Tabgha on the Sea of ​​Galilee , but without carrying out a detailed investigation. In 1909 the cancer Typhlocaris galilea, which only lives there, was discovered in this source together with Tethysbaena relicta , both of whom have a close relative in the Ajalon cave. Further research in the cave of Movile and the caves of Frasassi did not lead to the realization until the 1990s that subterranean ecosystems based on chemoautotrophy can exist.

Bacteria, such as those of the genus Beggiatoa , gain energy from the hydrogen sulfide in the water, which they use to form biomass from the carbon dioxide dissolved in the water . Hydrogen sulfide and sulfides are toxic to aerobic organisms and cause hypoxia by binding oxygen . An environment rich in sulfur compounds therefore requires special adaptations from higher organisms. Such adaptations are, for example, the formation of oxygen-binding proteins for transport or storage in the body, or endosymbiosis with sulfur-oxidizing bacteria .

biodiversity

Caves are usually poor in species compared to above-ground habitats. In a few caves, however, it was found that they harbor a comparatively wide range of species. Factors that caves with high biodiversity often have in common are the location in a karst landscape, the reaching down into the phreatic zone , a high production of organic matter, in particular through chemoautotrophy , and a great length. The Ajalon Cave combines all of these factors. A comparative study of cave faunas based on chemoautotrophy showed that a higher proportion of chemoautotrophic biomass was associated with more complex communities and greater species diversity in invertebrates . The special significance of the Ajalon Cave is also due to the fact that a closed system based entirely on chemoautotrophy was found here to socialize water dwellers and land animals.

Shortly after its discovery, the Ajalon Cave was examined by employees of the Hebrew University of Jerusalem . They found various bacteria, protozoa, and four types of crustaceans in the salty water of the cave lake . The crustaceans living in the cave lake are partly of marine origin and partly related to freshwater crabs. Four land-bound invertebrates have been discovered on the dry part of the lower level, but always near the cave lake . Among them were Akrav israchanani , a blind scorpion , of which only about 20, according to other sources 32, dead specimens were found, and the pseudoscorpion Ayyalonia dimentmani . Six of the animal species were previously unknown and were described for the first time in the following years, with a springtail of the genus Troglopedetes the identification at the species level and, if necessary, the first description are still pending .

Macrofauna

Macrofauna of the Ajalon cave
Scientific name Class, okay Endemic Remarks
Akrav israchanani Arachnids , scorpions ( Scorpiones ) Yes extinct, only about 20 dried up casings in the collection of the Hebrew University of Jerusalem, according to Israel Naaman remains of 32 dead animals found; the classification into a new monotypical family has been questioned
Ayyalonia dimentmani Arachnids , pseudoscorpions ( Pseudoscorpiones ) Yes often, on the rocks around the cave lake
Lepidospora ayyalonica Insects , fish ( zygentoma ) Yes only a male specimen, possibly only entered the cave after opening
Troglopedetes sp. Springtails , Entomobryomorpha (Yes) probably a new species; Identification at species level not yet possible due to lack of comparative material
Tethysbaena ophelicola Higher crustaceans , Thermosbaenacea Yes marine origin, all life stages in large quantities in the cave lake
Typhlocaris ayyaloni Higher crabs , decapods ( Decapoda ) Yes marine origin, hundreds in the cave lake, but no youth stages, the largest living beings in the Ajalon cave
Metacyclops longimaxillis Copepods , Cyclopoida Yes in comparison with other species of the genus, the maxillae are remarkably long ; in the Ajalon Cave in very large numbers of all ages
Metacyclops subdolus auctorum Copepods , Cyclopoida No since 1938 finds in Italy and other European Mediterranean countries, in sources at the Dead Sea and in the northern Negev desert; only a few adult and young animals in the Ajalon cave
Akrav israchanani , dried up shell
Typhlocaris ayyaloni , the largest creature in the cave
Tethysbaena ophelicola , female with brood pouches

The distribution of troglobiont scorpions is largely restricted to the tropics. Finding a type of cave-dwelling scorpion in Israel, outside the tropics, was therefore surprising. Together with the crustaceans in the cave, they were interpreted as relic fauna from the time of the tropical ocean Tethys . They see other attempts at explanation as part of an underground ecosystem that has developed independently. Finally, the possibility is considered that the scorpions did not evolve with the crustaceans, but rather colonized and trapped the cave at a later date.

There are closely related species in Israel of the crustaceans Typhlocaris ayyaloni and Tethysbaena ophelicola that live in the cave lake . Typhlocaris galilea comes from a single site on the Sea of ​​Galilee . Also Tethysbaena relicta is only localities in the flow of groundwater beneath the Jordan Valley known. This groundwater system has no direct connection to the Yarkon Taninim Aquifer and the Ajalon Cave. It is believed that both types of Ajalon Cave originated in the Jordan Rift and that they were isolated a long time ago. The two species of the genus Metacyclops are represented in very different numbers in the cave lake. From Metacyclops subdolus few adult and young animals were found during Metacyclops longimaxillis present in very large numbers in all age groups. Metacyclops longimaxillis seems to be better adapted to the high temperature and the high salt and sulfur content of the cave lake and therefore to multiply.

In order to determine the extent of the underground ecosystem, investigations were also carried out outside the cave. For this purpose, water from boreholes that are set in the quarry for safety reasons and extend into the groundwater, old observation wells of the water authority, abandoned wells and pools within a radius of several hundred meters were checked. Metacyclops subdolus was found in boreholes near the cave and in an above ground pool fed by groundwater. The same applies to Typhlocaris ayyaloni , of which hundreds of adult animals were found in the cave lake, but no juveniles or egg-bearing females. Animals of this species were also caught in boreholes outside the cave. It is assumed that these species enter the cave with inflowing groundwater or actively seek them out for food, and that they reproduce outside the cave under more favorable water conditions.

For other habitats of endemic crustaceans in Israel, such as the El Nur spring in Tabgha where Typhlocaris galilea was found , the occurrence of several species of snails , roundworms and a little bristle has been described. There are no representatives of these animal groups in the Ajalon Cave, but burial tunnels were discovered in the clay soil of an originally water-filled part of the cave, which cannot be associated with any of the animal species found.

Microfauna

The first research on chemolithoautotrophic bacteria from sulfur sources was carried out by Sergei Nikolajewitsch Winogradski in the 1880s . During the following century, research into such microorganisms was made difficult by the fact that they often cannot be distinguished microscopically and that most chemoautotrophic bacteria cannot be cultivated in the laboratory. Only genetic analysis made it possible to precisely identify the bacteria found, and isotope examinations of bacterial accumulations and samples of air, water and rocks helped to illustrate the metabolic processes. It was found that the bacterial faunas of the caves investigated so far are very complex; they all have in common the presence of green sulfur bacteria , gammaproteobacteria and epsilonproteobacteria . An exhaustive investigation of the entire bacterial fauna and the material cycles associated with it has not yet taken place in any cave.

The Ajalon Cave's ecosystem is based on the biomass produced by large quantities of sulfur-oxidizing bacteria . Mats made of bacteria float on 40 to 100% of the surface of the cave lake, and the shores of the lake are covered by them. The bacterial lawns consist primarily of Beggiatoa -like, thread-like bacteria with sulfur trapped in their vacuoles . Calcite crystals form on the bacterial mats, and if the mats exceed a critical mass as a result, they sink to the bottom of the lake. Since no remains of these mats were found on the bottom of the lake, it is unclear whether the mats reappear later or whether calcite and bacterial mats dissolve in the deeper water layers. In addition, other bacteria and, as single-celled organisms, numerous ciliates and amoebozoa were found. Until 2013, neither the bacterial fauna nor the unicellular organisms of the Ajalon cave had been examined in detail.

Tethysbaena ophelicola , subadult animals, the intestine filled with bacteria is highlighted

Food chain

During the isotope investigation of the scorpions in the Ajalon Cave, a PDB value of about –0.36 ‰ was measured. Feeding soil organisms from a normal atmosphere would result in a value of -0.25 to -0.18 ‰, and the deviating values ​​indicate feeding on organic matter from the cave. The isotope ratios of oxygen and carbon in bacteria and higher organisms in the cave showed that the bacteria are the energy source of the entire ecosystem.

The examined intestines of the particularly numerous crustaceans of the species Tethysbaena ophelicola were literally crammed with bacterial cells. An examination of the intestinal contents of two specimens of the species Typhlocaris ayyaloni showed that they too ate directly from the bacterial lawn and from smaller crustaceans of the species Tethysbaena ophelicola . It has not been clarified whether it is simply a matter of food utilization or whether the crustaceans maintain an endosymbiotic relationship with the bacteria .

There are various attempts to explain the further development of the food chain or the aquatic and terrestrial food chains. One approach provides that the very large numbers of Metacyclops longimaxillis and Tethysbaena ophelicola are permanent residents of the cave lake and, as consumers of bacteria, are at the beginning of the food chain. Typhlocaris ayyaloni and Metacyclops subdolus , on the other hand, have their actual habitat in other areas of the groundwater and only visit the cave lake to eat. As far as stygobionts are concerned, Typhlocaris ayyaloni is without a doubt at the end of the food chain.

In relation to the terrestrial inhabitants of the cave, the springtail Troglopedetes sp. regarded as the primary consumer who feeds directly on the bacteria on the shore of the cave lake and was also found on the bacteria mats floating in the water. It in turn serves the predatory pseudoscorpions as food. A study of the biology and ecology of Akrav israchanani was no longer possible, but other cave-dwelling scorpions are the most important predators in their ecosystems .

Biotope and species protection

Groundwater lowering

Ayyalonia dimentmani , a pseudoscorpion from the Ayalon cave

The rarity of such ecosystems as the Ajalon Cave, their high degree of biodiversity and the high proportion of endemics in their fauna has already led to the demand for immediate measures to be taken to protect them. The accidental opening of an entrance to the cave may have affected the biome . The lowering of the groundwater level due to excessive water abstraction from the aquifer has a much stronger impact . In the area of ​​the Ajalon Cave, a drop in the groundwater level of 13 meters has been determined since 1951. This sank the area of ​​the cave lake from around 4,000 to around 400 square meters, and a large part of the passages and chambers still in the process of cave formation fell dry.

The lower production of biomass due to the reduction in the size of the habitat is seen as a possible reason for the extinction of the scorpion Akrav israchanani , which, with its position at the end of the food chain, was particularly affected by disturbances in the ecosystem. This also indicates that most of the dead scorpions were found on the cave walls a few meters above the current water level. The location of the dead scorpions and a comparison with the reconstructed water levels in the cave made it possible to establish that Akrav israchanani became extinct between 1960 and 1991. Some arachnologists counter the hypotheses of a creeping extinction that scorpions react to a lack of food with cannibalism, for which there is no evidence of the dead scorpions found. They consider a sudden catastrophic event such as the eruption of large quantities of hydrogen sulfide as an explanation, but without explaining the survival of the pseudoscorpions and springtails in the cave.

In October 2010 the groundwater level in the area of ​​the Ajalon Cave reached a historic low of around 11.30 meters above sea level. This means that the cave lake is about to fall dry, there is no longer any space for the bacterial mats on the water surface. Even if there is still chemoautotrophic energy generation by bacteria on the underground contact surfaces of the thermal water with the groundwater, there is an immediate danger of extinction for the land animals of the Ajalon cave.

Quarry

Immediately after the discovery of the Ajalon Cave, Nesher Israel Cement Enterprises, as the operator of the quarry, and the Israeli Ministry of Infrastructure took joint measures to secure and maintain the cave. The operator of the quarry stated that regardless of possible disruptions to operations, his company is interested in preserving the cave and its ecosystem. To secure the cave as a natural monument, a trapezoidal area in the quarry is to remain untouched in the area of ​​the cave, while the planned mining continues around it.

Invasion of foreign species

Since the cave's ecosystem is in great danger from outside animals, access was secured early with a door. Its edges and the connections between the cave and the outside world that were created during earlier exploratory drilling were filled with polyurethane foam . This was also necessary because spiders from the aboveground fauna had already been caught in the cave, which had apparently entered the cave through a borehole. Despite such efforts and the commitment to preserving the Ajalon Cave as a karst phenomenon, concerns about the threat to the ecosystem persist. Half of the rock layer above the cave has already been removed and cracks may form in the rock due to the mining operations around the cave. This increases the risk of penetration by aboveground organisms, which could further disrupt the sensitive system and destroy individual fauna elements.

In the Red List of Threatened Species of the IUCN , the species Typhlocaris ayyaloni is classified as "Endangered" ( EN ). The classification is justified with the small number of sites and the observed deterioration of the habitat . For the other species of the Ajalon cave, no classification was made until 2013.

The Israeli Nature Conservation Act of 1998 (Article 33 (a) of Act 5758-1998) empowers the Minister for the Environment to enact an ordinance on Protected Natural Assets , which are also protected outside of designated nature reserves . The 2005 nature conservation ordinance ( Declaration on National Parks, Nature Reserves, National Sites and Memorial Sites Proclamation (Protected Natural Assets), 5765-2005 ) names numerous animal and plant species, fossils and geological formations and is based on the Israeli red for vertebrates and plants Lists . Of the species in the Ajalon cave, only the genus Typhlocaris is listed by name.

The "Ophel" as a global biome

From the assumption that individual species of the Ajalon cave only visit the cave lake with its rich food resources for food intake, it follows that there is a transport of biomass from the cave lake into the surrounding groundwater body. This possible horizontal transport of energy and the worldwide spread of crustaceans of the order Thermosbaenacea , which is represented in the Ajalon cave with Tethysbaena ophelicola , forms the basis of the theory of a global, underground and of the entry of energy developed by the zoologist Francis Dov Por externally independent biome, which he calls "Ophel" . In the “Ophel”, sulfur bacteria and other chemoautotrophic bacteria form biomass from sulfur-containing thermal water, and higher organisms feed on the bacteria. Por's theory did not meet with unqualified support. The Romanian zoologist Ştefan Negrea pointed out that there can be no completely isolated systems in nature.

Media reactions and publications

On May 31, 2006, the discovery of the Ajalon Cave and its extraordinary fauna were described by the zoologists Amos Frumkin and Chanan Dimentman at a press conference at the Hebrew University of Jerusalem. This was followed by extensive coverage in Israeli television programs and in major newspapers such as Haaretz , Ma'ariv , Jedi'ot Acharonot and the Jerusalem Post . International media also initially reported the discovery, but in the years that followed, the Ajalon Cave was only occasionally the subject of reports in Israel.

The first scientific publication took place in the journal Nature on June 8, 2006 under the short news. Francis Dov Por 2007 provided a detailed description of the findings available up to that point . In his publication he presented the ecosystem of the Ajalon cave based on chemolithoautotrophy and defined the Ophel as the second underground biosphere. With his diploma thesis from 2011, Israel Naaman provided the most comprehensive presentation to date of the formation of the Ajalon Cave and its influence by the anthropogenic sinking of the water table over the past decades. The processing of the zoological finds, including the first description of the newly discovered species and their publication, has not yet been fully completed. The status of two metazoa is still unclear and no detailed information is available on the microflora. In professional circles, especially among speleobiologists, the Ajalon cave with its fauna is still of great interest and it is mentioned again and again in scientific publications on speleology .

literature

Web links

Commons : Ajalon Cave  - Collection of pictures, videos and audio files

Individual evidence

  1. ↑ a b Boaz Langford, Amos Frumkin: The longest limestone caves of Israel , p. 106.
  2. ↑ Moshe Tsurnamal: A new species of the stygobiotic blind prawn Typhlocaris Calman, 1909 (Decapoda, Palaemonidae, Typhlocaridinae) from Israel , S. 490th
  3. ↑ a b c d e Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 1.
  4. ↑ a b Tamara Traubman: חיו בבועה, עד שהחוקרים הגיעו לרמלה. חור במחצבה חשף מערה קדומה ובה מינים לא מוכרים של סרטנים, עקרבים וחרקים (German: "Life in a bubble until researchers came to Ramla. In the quarry cave with unknown crabs, scorpions discovered") . In: Haaretz , June 1, 2006, online (accessed March 11, 2014).
  5. ↑ a b Moshe Tsurnamal: A new species of the stygobiotic blind prawn Typhlocaris Calman, 1909 (Decapoda, Palaemonidae, Typhlocaridinae) from Israel , S. 488th
  6. ↑ Danielle Defaye, Francis Dov Por: Metacyclops (Copepoda, Cyclopidae) from Ayyalon Cave, Israel , p. 401.
  7. ↑ Amos Frumkin: Active hypogene speleogenesis and groundwater system at the edge of an anticlinal ridge , p. 139.
  8. ↑ Tamara Traubman: Quarry cave lost in time yields' unknown species . In: Haaretz , June 1, 2006, online (accessed March 15, 2014).
  9. ^ Boaz Langford, Amos Frumkin: The longest limestone caves of Israel , p. 105.
  10. ↑ a b Amos Frumkin: Active hypogene speleogenesis and groundwater system at the edge of an anticlinal ridge , p. 142.
  11. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , pp. 23-25.
  12. ↑ a b c Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 26.
  13. ↑ a b c d Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 27.
  14. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , p. 28.
  15. ↑ a b c d Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 37.
  16. ↑ a b Yossef H. Hatzor, Ilia Wainshtein, Dagan Bakun Mazor: Stability of shallow karstic caverns in blocky rock masses , p. 1297.
  17. ↑ a b Yossef H. Hatzor, Ilia Wainshtein, Dagan Bakun Mazor: Stability of shallow karstic caverns in blocky rock masses , p. 1299.
  18. ↑ a b c Israel Naaman, Chanan Dimentman, Amos Frumkin: Active Hypogene Speleogenesis in a Regional Karst Aquifer: Ayyalon Cave, Israel , p. 73.
  19. ↑ a b c Ştefan Negrea: A remarkable finding that suggests the existence of a new groundwater biome based on chemoautotrophic resources, named "Ophel" by FD Por , p. 85.
  20. ↑ a b Gershom Levy: The first troglobite scorpion from Israel and a new chactoid family (Arachnida: Scorpiones) , p. 91.
  21. ↑ a b c H. P. Wagner: Tethysbaena ophelicola n. Sp. (Thermosbaenacea), a new prime consumer in the Ophel biome of the Ayyalon Cave, Israel , p. 1572.
  22. ↑ a b c Francis Dov Por: Groundwater life: some new biospeleological views resulting from the Ophel paradigm , p. 63.
  23. ↑ United Nations Economic and Social Commission for Western Asia, Federal Institute for Geosciences and Raw Materials (Ed.): Inventory of Shared Water Resources in Western Asia , p. 466.
  24. ↑ a b c d Shoshana Gabbay: The Environment in Israel , State of Israel, Ministry of the Environment, Jerusalem 2002, pp. 76-77, Online PDF, 20.2 MBhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fwww.sviva.gov.il%2FEnglish%2FResourcesandServices%2FPublications%2FDocuments%2FTheEnvironmentInIsrael2002.pdf~GB%3D~IA~3D~M0DZ .3D%3D SZ% 3D ~ double-sided% 3D ~ LT% 3DOnline% 20PDF% 2C% 2020% 2C2% 26nbsp% 3BMB ~ PUR% 3D (accessed March 20, 2014).
  25. ↑ a b Amos Frumkin: Active hypogene speleogenesis and groundwater system at the edge of an anticlinal ridge , p. 138.
  26. ↑ United Nations Economic and Social Commission for Western Asia, Federal Institute for Geosciences and Raw Materials (Ed.): Inventory of Shared Water Resources in Western Asia , p. 468.
  27. ↑ Amos Frumkin: Active hypogene speleogenesis and groundwater system at the edge of an anticlinal ridge , p. 137.
  28. ↑ United Nations Economic and Social Commission for Western Asia, Federal Institute for Geosciences and Raw Materials (Ed.): Inventory of Shared Water Resources in Western Asia , pp. 473–474.
  29. ↑ United Nations Economic and Social Commission for Western Asia, Federal Institute for Geosciences and Raw Materials (Ed.): Inventory of Shared Water Resources in Western Asia , pp. 474–475.
  30. ↑ a b Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 3.
  31. ↑ Amos Frumkin, Haim Gvirtzman: Cross-formational rising groundwater at an artesian karstic basin: the Ayalon Saline Anomaly, Israel , pp. 317-318.
  32. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , p. 5.
  33. ↑ a b Amos Frumkin, Haim Gvirtzman: Cross-formational rising groundwater at an artesian karstic basin: the Ayalon Saline Anomaly, Israel , p. 331.
  34. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , pp. 69–70.
  35. ↑ a b c Israel Naaman, Chanan Dimentman, Amos Frumkin: Active Hypogene Speleogenesis in a Regional Karst Aquifer: Ayyalon Cave, Israel , pp. 73-74.
  36. ↑ a b Jennifer L. Macalady et al .: Dominant Microbial Populations in Limestone-Corroding Stream Biofilms, Frasassi Cave System, Italy . In: Applied and Environmental Microbiology , Volume 72, Number 8, pp. 5596-5609, here p. 5596, doi : 10.1128 / AEM.00715-06 .
  37. ^ Annette Summers Engel: Observations on the biodiversity of sulfidic karst habitats , pp. 188-189.
  38. ↑ a b Annette Summers Engel: Observations on the biodiversity of sulfidic karst habitats , p. 192.
  39. ↑ a b c Božidar PM Ćurčić: Ayyalonia dimentmani ng, n. Sp. (Ayyaloniini n. Trib., Chthoniidae, Pseudoscorpiones) from a cave in Israel . In: Archives of Biological Sciences , Volume 60, Number 3, pp. 331-339, here p. 332, doi : 10.2298 / ABS0803331C .
  40. ↑ Victor Fet, Michael E. Soleglad, Sergei L. Zonstein: The Genus Akrav Levy, 2007 (Scorpiones: Akravidae) Revisited , p. 5.
  41. ↑ a b Moshe Tsurnamal: A new species of the stygobiotic blind prawn Typhlocaris Calman, 1909 (Decapoda, Palaemonidae, Typhlocaridinae) from Israel , S. 498th
  42. ^ A b c Francis Dov Por: Groundwater life: some new biospeleological views resulting from the Ophel paradigm , p. 62.
  43. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , p. 14.
  44. ^ A b c Francis Dov Por: Ophel, the Newly Discovered Hypoxic Chemolithotrophic Groundwater Biome: A Window to Ancient Animal Life , p. 467.
  45. ↑ Annette Summers Engel: Observations on the biodiversity of sulfidic karst habitats , pp. 187–188.
  46. ↑ a b Jean-François Flot, Gert Wörheide, Sharmishtha Dattagupta: Unsuspected diversity of Niphargus amphipods in the chemoautotrophic cave ecosystem of Frasassi, central Italy . In: BMC Evolutionary Biology , Volume 10, 2010, Article 171, pp. 1-2, doi : 10.1186 / 1471-2148-10-171 .
  47. ↑ Stéphane Hourdez, François H. Lallier: Adaptations to hypoxia in hydrothermal-vent and cold-seep invertebrates , pp. 143-144.
  48. ↑ Manfred K. Grieshaber, Susanne Völkel: Animal adaptations for tolerance and exploitation of poisonous sulfide , p. 36.
  49. ↑ a b Manfred K. Grieshaber, Susanne Völkel: Animal adaptations for tolerance and exploitation of poisonous sulfide , pp. 36-42.
  50. ↑ Stéphane Hourdez, François H. Lallier: Adaptations to hypoxia in hydrothermal-vent and cold-seep invertebrates , pp. 144-145.
  51. ↑ Stéphane Hourdez, François H. Lallier: Adaptations to hypoxia in hydrothermal-vent and cold-seep invertebrates , pp. 151–153.
  52. ↑ a b c David C. Culver, Boris Sket: Hotspots of Subterranean Biodiversity in Caves and Wells . In: Journal of Cave and Karst Studies , Volume 62, Number 1, Pages 11–17, here p. 16, Online PDF, 75 kBhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fmail.caves.org%2Fpub%2Fjournal%2FPDF%2FV62%2Fv62n1-Culver.pdf~GB%3D~IA%3D~MDZ%3D%0A~ SZ% 3D ~ double-sided% 3D ~ LT% 3DOnline% 20PDF% 2C% 2075% 26nbsp% 3BkB ~ PUR% 3D (accessed on March 21, 2014).
  53. ↑ Megan L. Porter et al .: Productivity-Diversity Relationships from Chemolithoautotrophically Based Sulfidic Karst Systems . In: International Journal of Speleology , Volume 38, Number 1, 2009, pp. 27-40.
  54. ↑ Annette Summers Engel: Observations on the biodiversity of sulfidic karst habitats , pp. 195–198.
  55. ↑ a b c d Francis Dov Por et al .: Animal life in the chemoautotrophic ecosystem of the hypogenic groundwater cave of Ayyalon (Israel) , p. 10.
  56. ↑ a b without author: Aquatic Invertebrates, with the Arachnid and the Medical Parasitological Collections . In: Haasiana , number 3, 2006, pp. 56-63, here p. 58, ISSN  0793-5862 .
  57. ↑ a b c d e Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 46.
  58. ↑ a b Victor Fet, Michael E. Soleglad, Sergei L. Zonstein: The Genus Akrav Levy, 2007 (Scorpiones: Akravidae) Revisited , p. 4.
  59. ↑ a b c d e Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 47.
  60. ^ Francis Dov Por et al .: Animal life in the chemoautotrophic ecosystem of the hypogenic groundwater cave of Ayyalon (Israel) , pp. 9-10.
  61. ↑ Luis F. Mendes et al .: New data and new species of Microcoryphia and Zygentoma (Insecta) from Israel . In: Annales de la Société Entomologique de France (ns) , Volume 47, Number 3-4, 2011, pp. 384-393, here pp. 392-393, doi : 10.1080 / 00379271.2011.10697732 .
  62. ↑ a b c H. P. Wagner: Tethysbaena ophelicola n. Sp. (Thermosbaenacea), a new prime consumer in the Ophel biome of the Ayyalon Cave, Israel , p. 1574.
  63. ↑ Danielle Defaye, Francis Dov Por: Metacyclops (Copepoda, Cyclopidae) from Ayyalon Cave, Israel , p. 400.
  64. ↑ a b c d e Danielle Defaye, Francis Dov Por: Metacyclops (Copepoda, Cyclopidae) from Ayyalon Cave, Israel , p. 420.
  65. ↑ Danielle Defaye, Francis Dov Por: Metacyclops (Copepoda, Cyclopidae) from Ayyalon Cave, Israel , pp. 412-413.
  66. ^ Francis Dov Por et al .: Animal life in the chemoautotrophic ecosystem of the hypogenic groundwater cave of Ayyalon (Israel) , p. 9.
  67. ^ Wilson R. Lourenço, Renner Luiz Cerqueira Baptista, Alessandro Ponce de Leão Giupponi: Troglobitic scorpions: a new genus and species from Brazil . In: Comptes Rendus Biologies , Volume 327, Number 12, 2004, pp. 1151–1156, here p. 1153, doi : 10.1016 / j.crvi.2004.09.001 , Online PDF, 300 kBhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fwww.museunacional.ufrj.br%2Fmndi%2FAracnologia%2Faracnopdfs%2FLourenco%2520et%2520al%25202004%2520Troglo.pdf~GB3D MDZ% ​​3D% 0A ~ SZ% 3D ~ double-sided% 3D ~ LT% 3DOnline% 20PDF% 2C% 20300% 26nbsp% 3BkB ~ PUR% 3D (accessed on March 5, 2014) .
  68. ↑ a b Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 73.
  69. ↑ a b c d Gershom Levy: The first troglobite scorpion from Israel and a new chactoid family (Arachnida: Scorpiones) , p. 92.
  70. ↑ HP Wagner: Tethysbaena ophelicola n. Sp. (Thermosbaenacea), a new prime consumer in the Ophel biome of the Ayyalon Cave, Israel , pp. 1584-1586.
  71. ↑ a b Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 21.
  72. ↑ a b c Francis Dov Por: Groundwater life: some new biospeleological views resulting from the Ophel paradigm , p. 66.
  73. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , p. 69.
  74. ↑ Moshe Tsurnamal, Francis Dov Por: The subterranean fauna associated with the blind Palaemonid prawn Typhlocaris galilea Calman , pp. 220-221.
  75. ↑ a b Moshe Tsurnamal, Francis Dov Por: The subterranean fauna associated with the blind Palaemonid prawn Typhlocaris galilea Calman , S. 221st
  76. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , pp. 47–48.
  77. ↑ a b c d Annette Summers Engel: Observations on the biodiversity of sulfidic karst habitats , p. 190.
  78. ↑ Annette Summers Engel: Observations on the biodiversity of sulfidic karst habitats , pp. 190–191.
  79. ↑ a b c Israel Naaman, Chanan Dimentman, Amos Frumkin: Active Hypogene Speleogenesis in a Regional Karst Aquifer: Ayyalon Cave, Israel , p. 74.
  80. ↑ a b c d Francis Dov Por et al .: Animal life in the chemoautotrophic ecosystem of the hypogenic groundwater cave of Ayyalon (Israel) , p. 7.
  81. ↑ Victor Fet, Michael E. Soleglad, Sergei L. Zonstein: The Genus Akrav Levy, 2007 (Scorpiones: Akravidae) Revisited , p. 46.
  82. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , p. 68.
  83. ↑ Moshe Tsurnamal: A new species of the stygobiotic blind prawn Typhlocaris Calman, 1909 (Decapoda, Palaemonidae, Typhlocaridinae) from Israel , S. 499th
  84. ^ Francis Dov Por: Groundwater life: some new biospeleological views resulting from the Ophel paradigm , p. 64.
  85. ^ Francis Dov Por: Groundwater life: some new biospeleological views resulting from the Ophel paradigm , p. 67.
  86. ↑ James R. Reddell: Spiders and related groups In: William B. White, David C. Culver (Eds.): Encyclopedia of Caves. Second Edition , Academic Press, Waltham, MA 2012, pp. 786-797, here p. 787, ISBN 978-0-12-383832-2 .
  87. ↑ Victor Fet: Заметки о скорпионах и скорпиологах (German: "Notes on Scorpions"). In: Природа (German: “Priroda” / “Natur”), number 10, 2013, pp. 52–58, ISSN  0032-874X , online PDF, 726 kBhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fwww.science.marshall.edu%2Ffet%2Feuscorpius%2FFet_Priroda%25202013_10.pdf~GB%3D~IA%3D~MDZ%3D%0A~SZ% 3D ~ double-sided% 3D ~ LT% 3DOnline% 20PDF% 2C% 20726% 26nbsp% 3BkB ~ PUR% 3D (Russian, accessed on March 15, 2014), on p. 56 detailed photos of Akrav israchanani with the hook-shaped tips of the pliers.
  88. ↑ HP Wagner: Tethysbaena ophelicola n. Sp. (Thermosbaenacea), a new prime consumer in the Ophel biome of the Ayyalon Cave, Israel , pp. 1572-1574.
  89. ↑ a b Ofri Ilani: One year later, 'Noah's Ark' cave is no longer a safe haven . In: Haaretz , July 19, 2007, online (accessed March 15, 2014).
  90. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , pp. 70–71.
  91. ↑ a b Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 71.
  92. ↑ Victor Fet, Michael E. Soleglad, Sergei L. Zonstein: The Genus Akrav Levy, 2007 (Scorpiones: Akravidae) Revisited , pp. 6-7.
  93. ↑ Yossef H. Hatzor, Ilia Wainshtein, Dagan Bakun Mazor: Stability of shallow karstic caverns in blocky rock masses , pp. 1298-1299.
  94. ↑ a b Israel Naaman: Karst system and ecology of the Ayalon cave, Israel , p. 20.
  95. ↑ Israel Naaman: Karst System and Ecology of the Ayalon Cave, Israel , pp. 72–73.
  96. ↑ Typhlocaris ayyaloni in the IUCN Red List of Threatened Species 2013.2. Posted by: S. De Grave, 2012. Retrieved March 9, 2014 ..
  97. ↑ a b Richard Laster, Dan Livney: Environmental Law in Israel , Kluwer Law International, Alphen aan den Rijn 2011, p. 121, ISBN 978-90-4113610-7 .
  98. ↑ a b without author: Declaration on National Parks, Nature Reserves, National Sites and Memorial Sites Proclamation (Protected Natural Assets), 5765-2005 , Online PDF, 71 kBhttp: //vorlage_digitalisat.test/1%3Dhttp%3A%2F%2Fwww.sviva.gov.il%2FEnglish%2FLegislation%2FDocuments%2FNational%2520Parks%2C%2520Nature%2520Reserves%2C%2520National%2520Sites%2C% 2520Sites% 2520Laws% 2520and% 2520Regulations% 2FNationalParksNatureReservesNationalSitesAndMemorialSitesProclamation-2005.pdf ~ GB% 3D ~ IA% 3D ~ MDZ% 3D% 0A ~ SZ% 3D ~ double-sided% 3D ~ LT% 3DOnline% 20% 371% 2CBb% 2CB PUR% 3D (accessed on March 18, 2014).
  99. ^ A b Francis Dov Por et al .: Animal life in the chemoautotrophic ecosystem of the hypogenic groundwater cave of Ayyalon (Israel) , pp. 10-11.
  100. ↑ Ştefan Negrea: A remarkable finding that suggests the existence of a new groundwater biome based on chemoautotrophic resources, named "Ophel" by FD Por , p. 86.
  101. ↑ Without author: Unique Underground Ecosystem Revealed by Hebrew University Researchers Uncovers Eight Previously Unknown Species , press release of the Hebrew University of Jerusalem of May 31, 2006, online , accessed on March 19, 2014.
  102. ↑ a b without author: Aquatic Invertebrates, with the Arachnid and the Medical Parasitological Collections , p. 61.
  103. ↑ without author: Israeli cave reveals eight arthropod species (section “News in Brief”). In: Nature , volume 441, number 7094, p. 680, doi : 10.1038 / 441707a .