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endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found els ...
of Dinaric Alps. Subterranean fauna refers to Animal, animal species that are adaptation, adapted to live in an underground Natural environment, environment. Troglofauna and stygofauna are the two types of subterranean fauna. Both are associated with hypogean habitats – troglofauna is associated with terrestrial subterranean environment (
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 underground spaces above the water table), and stygofauna with all kind of subterranean waters (
groundwater Groundwater is the water present beneath Earth's surface in rock and soil pore spaces and in the fractures of rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit of rock or an unconsolidat ...
,
aquifer An aquifer is an underground layer of water-bearing, permeable rock, rock fractures, or unconsolidated materials ( gravel, sand, or silt). Groundwater from aquifers can be extracted using a water well. Aquifers vary greatly in their characteri ...
s, subterranean rivers, dripping bowls,
gours Rimstone, also called gours, is a type of speleothem (cave formation) in the form of a stone dam. Rimstone is made up of calcite and other minerals that build up in cave pools. The formation created, which looks like stairs, often extends into fl ...
, etc.).


Environment

Subterranean fauna is found worldwide and includes representatives of many
animal group In the English language, animals have different names depending on whether they are male, female, young, domesticated, or in groups. The best-known source of many English words used for collective groupings of animals is ''The Book of Saint Al ...
s, mostly
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s and other
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s. However, there is a number of
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
s (such as cavefishes and cave salamanders), although they are less common. Because of the complexity in exploring underground environments, many subterranean species are yet to be discovered and described. Peculiarities of underground habitat make it an extreme environment and, consequently, underground species are usually less than species living in epigean habitats. The main characteristic of underground environment is the lack of sunlight. Climatic values, like
temperature Temperature is a physical quantity that expresses quantitatively the perceptions of hotness and coldness. Temperature is measured with a thermometer. Thermometers are calibrated in various temperature scales that historically have relied on ...
and
relative humidity Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation, dew, or fog to be present. Humidity dep ...
, are generally almost stable – temperature corresponds to annual mean temperature in the place where the cavity opens, relative humidity rarely drops below 90%. Food sources are limited and localized. The lack of sunlight inhibits photosynthetic processes, so food comes only from epigean environment (through percolating water,
gravity In physics, gravity () is a fundamental interaction which causes mutual attraction between all things with mass or energy. Gravity is, by far, the weakest of the four fundamental interactions, approximately 1038 times weaker than the str ...
, or passive transport by animals). Important food sources in underground habitat are animals being decomposed and bat
guano Guano (Spanish from qu, wanu) is the accumulated excrement of Seabird, seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant ...
, that creates large
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
communities in such caves.


Ecological classification

Cave dwelling animals show different levels of adaptations to underground environment. According to a recent classification, animals living in terrestrial subterranean habitats can be classified into 3 categories, based on their
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overl ...
: * troglobionts (or troglobites): species strongly bound to subterranean habitats; * troglophiles: species living both in subterranean and in epigean habitats. Troglophiles are also divided in ''eutroglophiles'' (epigean species able to maintain a permanent subterranean population) and ''subtroglophiles'' (species inclined to perpetually or temporarily inhabit a subterranean habitat, but intimately associated with epigean habitats for some functions); * trogloxenes: species only occurring sporadically in a hypogean habitat and unable to establish a subterranean population. Regarding stygofauna, the corresponding words stygobionts (or stygobites), stygophiles and stygoxenes are used.


Biology

The cave beetle '' Leptodirus hochenwartii'' (family Leiodidae).">Leiodidae.html" ;"title="Leptodirus hochenwartii'' (family Leiodidae">Leptodirus hochenwartii'' (family Leiodidae). Characteristics of underground environment caused cave dwelling animals to evolve a number of adaptations, both
morphological and Physiology">physiological Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemica ...
. Examples of morphological adaptations include depigmentation (loss of external pigmentation), a reduction of cuticle thickness and the often extreme decrease of eyesight culminating in anophthalmia (complete loss of eyes). Exceptions, however, are harvestmen (Opiliones) in New Zealand caves, which possess large, functional eyes, presumably because these spider-like chelicerates feed on cave-dwelling, light-emitting glowworm larvae '' Arachnocampa'' which they detect visually. Other adaptations include the development and elongation of
antennal Antennae ( antenna), sometimes referred to as "feelers", are paired appendages used for sensing in arthropods. Antennae are connected to the first one or two segments of the arthropod head. They vary widely in form but are always made of one o ...
and locomotory appendages, in order to better move around and respond to environmental stimuli. These structures are well endowed with
chemical A chemical substance is a form of matter having constant chemical composition and characteristic properties. Some references add that chemical substance cannot be separated into its constituent elements by physical separation methods, i.e., wit ...
, tactile and humidity receptors (such as Hamann's organ in the cave
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
''Leptodirus'' ''hochenwartii''). Physiological adaptations include slow
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run c ...
and reduced
energy In physics, energy (from Ancient Greek: ἐνέργεια, ''enérgeia'', “activity”) is the quantitative property that is transferred to a body or to a physical system, recognizable in the performance of work and in the form of hea ...
consumption, due to limited food supply and low energy efficiency. This is likely to be realized through reducing movements, erasing aggressive interactions, improving feeding capability and food usage efficiency, and through ectothermy. As a consequence, cave dwelling animals can resist without eating for long time, live more than comparable epigean species, reproduce late in their lifespan, and produce fewer and bigger
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
s.


Evolution and ecology

Subterranean fauna have evolved in isolation. Stratigraphic barriers, such as rock walls and layers, and fluvial barriers, such as rivers and streams, prevent or hinder the dispersal of these animals. Consequently, subterranean fauna habitat and food availability can be very disjunct and precludes the great range of observed diversity across landscapes.


Threats to subterranean fauna

Floodwaters can be detrimental to subterranean species, by dramatically changing the availability of habitat, food and connectivity to other habitats and oxygen. Many subterranean fauna are likely to be sensitive to changes in their environment and floods, which can accompany a drop in temperature, may adversely affect some animals. Humans also pose a threat to troglofauna. Mismanagement of contaminants (e.g. pesticides and sewage) may poison subterranean fauna communities and removal of habitat (e.g. rising/lowering of the watertable or various forms of mining) can also be a major threat.


See also

* Cave conservation * List of troglobites *
Speleology Speleology is the scientific study of caves and other karst features, as well as their make-up, structure, physical properties, history, life forms, and the processes by which they form ( speleogenesis) and change over time (speleomorphology ...
* Subterranean river * Trogloxene


References

{{reflist, 24em Cave animals Subterranea (geography) Animals by adaptation Animals by location Ecology