Biospeleology, also known as cave biology, is a branch of

biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary i ...

dedicated to the study of

organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...

s that live in

cave A cave or cavern is a natural void in the ground, specifically a space large enough for a human to enter. Caves often form by the weathering of rock and often extend deep underground. The word ''cave'' can refer to smaller openings such as sea ...

s and are collectively referred to as

troglofauna Troglofauna are small cave-dwelling animals that have adapted to their dark surroundings. Troglofauna and stygofauna are the two types of subterranean fauna (based on life-history). Both are associated with subterranean environments – troglofaun ...

Biospeleology as a science

History

The first documented mention of a cave organisms dates back to 1689, with the documentation of the

olm The olm or proteus (''Proteus anguinus'') is an aquatic salamander in the family Proteidae, the only exclusively cave-dwelling chordate species found in Europe. In contrast to most amphibians, it is entirely aquatic, eating, sleeping, and br ...

, a cave salamander. Discovered in a cave in

Slovenia Slovenia ( ; sl, Slovenija ), officially the Republic of Slovenia (Slovene: , abbr.: ''RS''), is a country in Central Europe. It is bordered by Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the southeast, an ...

, in the region of

Carniola Carniola ( sl, Kranjska; , german: Krain; it, Carniola; hu, Krajna) is a historical region that comprised parts of present-day Slovenia. Although as a whole it does not exist anymore, Slovenes living within the former borders of the region sti ...

, it was mistaken for a baby dragon and was recorded by

Johann Weikhard von Valvasor Johann Weikhard Freiherr von Valvasor or Johann Weichard Freiherr von Valvasor ( sl, Janez Vajkard Valvasor, ) or simply Valvasor (baptised on 28 May 1641 – September or October 1693) was a natural historian and polymath from Carniola, pr ...

in his work ''

The Glory of the Duchy of Carniola ''The Glory of the Duchy of Carniola'' (german: Die Ehre deß Hertzogthums Crain, sl, Slava vojvodine Kranjske) is an encyclopedia published in Nuremberg in 1689 by the polymath Johann Weikhard von Valvasor. It is the most important work on his ...

''. The first formal study on cave organisms was conducted on the blind cave beetle. Found in 1831 by Luka Čeč, an assistant to the lamplighter, when exploring the newly discovered inner portions of the

Postojna cave Postojna Cave ( sl, Postojnska jama; german: Adelsberger Grotte; it, Grotte di Postumia) is a long karst cave system near Postojna, southwestern Slovenia. It is the second-longest cave system in the country (following the Migovec System) as wel ...

system in southwestern

.Vrezec A. et al. (2007
Monitoring populacij izbranih ciljnih vrst hroščev (končno poročilo)
(Monitoring of selected populations of target beetle species).

Natura 2000 Natura 2000 is a network of nature protection areas in the territory of the European Union. It is made up of Special Areas of Conservation and Special Protection Areas designated under the Habitats Directive and the Birds Directive, respectively ...

report. "Narrow-necked" blind cave beetle
. Slovenian museum of natural history. Accessed 2009-03-16. The specimen was turned over to Ferdinand J. Schmidt, who described it in the paper ''Illyrisches Blatt'' (1832). He named it ''

Leptodirus Hochenwartii ''Leptodirus'' is a cave beetle in the family Leiodidae. The genus contains only the single species ''Leptodirus hochenwartii''. It is a true troglobite, endemic to Slovenian, Croatian and, partly, Italian caves. Biology and ecology ''Leptodi ...

'' after the donor, and also gave it the Slovene name ''drobnovratnik'' and the

German German(s) may refer to: * Germany (of or related to) **Germania (historical use) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizens of Germany, see also German nationality law **Ger ...

name ''Enghalskäfer'', both meaning "slender-necked (beetle)".Mader B. (2003)
Archduke Ludwig Salvator and ''Leptodirus hohenwarti'' from Postonjska jama
. ''Acta carsologica'' 32(2): 290-298. The article represents the first formal description of a cave animal (the

, described in 1768, wasn't recognized as a cave animal at the time). Subsequent research by Schmidt revealed further previously unknown cave inhabitants, which aroused considerable interest among natural historians. For this reason, the discovery of ''L. hochenwartii'' (along with the olm) is considered as the starting point of biospeleology as a scientific discipline.Polak S. (2005). "Importance of discovery of the first cave beetle ''Leptodirus hochenwartii'' Schmidt, 1832.

''Endins'' 28 (2005). Biospeleology was formalized as a science in 1907 by

Emil Racoviță Emil Gheorghe Racoviță (; 15 November 1868 – 19 November 1947) was a Romanian biologist, zoologist, speleologist, and Antarctic explorer. Together with Grigore Antipa, he was one of the most noted promoters of natural sciences in Romani ...

with his seminal work ''Essai sur les problèmes biospéologiques'' ("Essay on biospeleological problems”).

Subdivisions

Organisms Categories

Cave organisms fall into three basic classes:

=Troglobite

= Troglobites are obligatory ''cavernicoles'', specialized for cave life. Some can leave caves for short periods, and may complete parts of their life cycles above ground, but cannot live their entire lives outside of a cave environment. Examples include chemotrophic bacteria, some species of

flatworms The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegment ...

springtails Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ent ...

, and

cavefish Cavefish or cave fish is a generic term for fresh and brackish water fish adapted to life in caves and other underground habitats. Related terms are subterranean fish, troglomorphic fish, troglobitic fish, stygobitic fish, phreatic fish and hypog ...

=Troglophile

= Troglophiles can live part or all of their lives in caves, but can also complete a life cycle in appropriate environments on the surface. Examples include

cave cricket The orthopteran family Rhaphidophoridae of the suborder Ensifera has a worldwide distribution. Common names for these insects include cave wētā, cave crickets, camelback crickets, camel crickets, Hogan bugs, spider crickets (sometimes shortene ...

bats Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...

millipede Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...

pseudoscorpion Pseudoscorpions, also known as false scorpions or book scorpions, are small, scorpion-like arachnids belonging to the order Pseudoscorpiones, also known as Pseudoscorpionida or Chelonethida. Pseudoscorpions are generally beneficial to humans sin ...

s and spiders.

=Trogloxene

Trogloxene Trogloxenes or subtroglophiles, also called cave guests, are animal species which periodically live in underground habitats such as caves or at the very entrance, but cannot live exclusively in such habitats."Cave Plants and Animals". 9 December Am ...

s frequent caves, and may require caves for a portion of its life cycle, but must return to the surface (or a ''parahypogean'' zone) for at least some portion of its life.

Oilbird The oilbird (''Steatornis caripensis''), locally known as the , is a bird species found in the northern areas of South America including the Caribbean island of Trinidad. It is the only species in the genus ''Steatornis'', the family Steatornith ...

s and most Daddy longlegs are trogloxenes.

Environmental Categories

Cave environments fall into three general categories:

= Endogean

= Endogean environments are the parts of caves that are in communication with surface soils through cracks and rock seams, groundwater seepage, and root protrusion.

=Parahypogean

= Parahypogean environments are the threshold regions near cave mouths that extend to the last penetration of sunlight.

=Hypogean

= Hypogean or "true" cave environments. These can be in regular contact with the surface via wind and underground rivers, or the migration of animals, or can be almost entirely isolated. Deep hypogean environments can host autonomous ecologies whose primary source of energy is not sunlight, but chemical energy liberated from limestone and other minerals by

chemoautotrophic A Chemotroph is an organism that obtains energy by the oxidation of electron donors in their environments. These molecules can be organic ( chemoorganotrophs) or inorganic (chemolithotrophs). The chemotroph designation is in contrast to phototro ...

bacteria.

Notable biospeleologists

, co-founder of biospeleology *

Carl Eigenmann Carl Henry Eigenmann (March 9, 1863 – April 24, 1927) was a German-American ichthyologist of the late nineteenth and early twentieth centuries, who, along with his wife Rosa Smith Eigenmann, and his zoology students is credited with identifying ...

Louis Fage Jean-Louis Fage (30 September 1883, in Limoges – 1964, in Dijon) was a French marine biologist and arachnologist. A native of Limoges, he studied biology at the Sorbonne and in the laboratory at Saint-Vaast-la-Hougue. In 1906 he obtained his do ...

René Jeannel René Jeannel (23 March 1879 – 20 February 1965) was a French entomologist.Jean-Jacques Amigo, « Jeannel (René, Gabriel, Marie) », in Nouveau Dictionnaire de biographies roussillonnaises, vol. 3 Sciences de la Vie et de la Terre, Perpignan, ...

, co-founder of biospeleology *

Curt Kosswig Dr. Curt Kosswig (sometimes spelled "Koßwig") (30 October 1903, Berlin – 29 March 1982, Hamburg) was a German zoologist and geneticist who spent most of his career at the University of Istanbul (1937–1955) and Hamburg University (1955–1969) ...

Bibliography

* Bernard Collignon, Caving, scientific approaches., Edisud 1988 * Fabien Steak, Approach biospéologie, File EFS Instruction No. 116, 1st Edition, 1997 * C. Delamare-Debouteville, Life in caves, PUF, Que sais-je?, Paris 1971 * Bernard Gèze, Scientific caving, Seuil, Paris, 1965, p. 137-167 * R. and V. Decou Ginet, Introduction to biology and groundwater ecology, University Publishing Delarge 1977 * René Jeannel, Animal cave in France, Lechevalier, Paris, 1926 * René Jeannel, Living fossils caves, Gallimard, Paris, 1943 * Edward Alfred Martel, Groundwater evolution, Flammarion, Paris, 1908, p. 242-289 * Georges Émile Racovitza, Essay on biospéologiques problems, I Biospeologica 1907 * Michel Siffre, Animals sinkholes and caves, Hachette, 1979 * Michel Siffre, France The caves and caverns, ed. Privat, 1999, p. 136-153 * G. and R. Thines Tercafs, Atlas of the underground life: the cave animals, Boubée (Paris) and De Visscher (Brussels), 1972 * Albert Vandel Biospéologie: the biology of cave animals, Gauthier-Villars, Paris, 1964 * Armand Vire, The subterranean fauna of France, Paris, 1900

References

External links

Biospeleology; The Biology of Caves, Karst, and Groundwater
by Texas Natural Science Center, the

University of Texas at Austin The University of Texas at Austin (UT Austin, UT, or Texas) is a public research university in Austin, Texas. It was founded in 1883 and is the oldest institution in the University of Texas System. With 40,916 undergraduate students, 11,075 ...

and the

Missouri Department of Conservation The Missouri Department of Conservation (MDC) and the Missouri Conservation Commission were created by Article IV Sections 40-42 of the Missouri Constitution, which were adopted by the voters of the state in 1936 as Amendment 4 to the constitution ...

Cave Biota; an evolving "webumentary"
sponsored by

Hoosier National Forest The Hoosier National Forest is a property managed by the United States Forest Service in the hills of southern Indiana. Composed of four separate sections, it has a total area of . Hoosier National Forest's headquarters are located in Bedford, w ...

and

Indiana Karst Conservancy Indiana () is a U.S. state in the Midwestern United States. It is the 38th-largest by area and the 17th-most populous of the 50 States. Its capital and largest city is Indianapolis. Indiana was admitted to the United States as the 19th sta ...

{{Caves Speleology Earth sciences