The 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
Stygofauna are any fauna that live in groundwater systems or aquifers, such as caves, fissures and vugs. Stygofauna and troglofauna are the two types of subterranean fauna (based on life-history). Both are associated with subterranean environmen ...
are the two types of subterranean fauna. Both are associated with hypogeal habitats – troglofauna is associated with terrestrial subterranean environment (
cave
Caves or caverns are natural voids under the Earth's Planetary surface, surface. Caves often form by the weathering of rock and often extend deep underground. Exogene caves are smaller openings that extend a relatively short distance undergrou ...
s and underground spaces above the
water table
The water table is the upper surface of the phreatic zone or zone of saturation. The zone of saturation is where the pores and fractures of the ground are saturated with groundwater, which may be fresh, saline, or brackish, depending on the loc ...
), and stygofauna with all kind of subterranean waters (
groundwater
Groundwater is the water present beneath Earth's surface in rock and Pore space in soil, soil pore spaces and in the fractures of stratum, rock formations. About 30 percent of all readily available fresh water in the world is groundwater. A unit ...
,
aquifer
An aquifer is an underground layer of water-bearing material, consisting of permeability (Earth sciences), permeable or fractured rock, or of unconsolidated materials (gravel, sand, or silt). Aquifers vary greatly in their characteristics. The s ...
s,
subterranean river
A subterranean river (also known as an underground river) is a river or watercourse that runs wholly or partly beneath the ground, one where the riverbed does not represent the surface of the Earth. It is distinct from an aquifer, which may flow ...
s, dripping bowls,
gours, etc.).
Environment
Subterranean fauna is found worldwide and includes representatives of many
animal groups, mostly
arthropod
Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s and other
invertebrate
Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s. However, there is a number of
vertebrate
Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain.
The vertebrates make up the subphylum Vertebra ...
s (such as
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, Troglomorphism, troglomorphic fish, troglobitic fish, stygobitic fish, phreat ...
es and
cave salamander
A cave salamander is a type of salamander that primarily or exclusively inhabits caves, a group that includes several species. Some of these animals have developed special, even extreme, adaptations to their subterranean environments. Some specie ...
s), 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
Sunlight is the portion of the electromagnetic radiation which is emitted by the Sun (i.e. solar radiation) and received by the Earth, in particular the visible spectrum, visible light perceptible to the human eye as well as invisible infrare ...
. Climatic values, like
temperature
Temperature is a physical quantity that quantitatively expresses the attribute of hotness or coldness. Temperature is measurement, measured with a thermometer. It reflects the average kinetic energy of the vibrating and colliding atoms making ...
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 (meteorology), precipitation, dew, or fog t ...
, 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
Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin and contains essential nutrients such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is inges ...
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 (), also known as gravitation or a gravitational interaction, is a fundamental interaction, a mutual attraction between all massive particles. On Earth, gravity takes a slightly different meaning: the observed force b ...
, or passive transport by animals). An exception are caves like the
Movile Cave
Movile Cave () is a cave near Mangalia, Constanța County, Romania discovered in 1986 by Cristian Lascu during construction work a few kilometers from the Black Sea coast. It is notable for its unique subterranean groundwater ecosystem abund ...
, where
chemosynthesis
In biochemistry, chemosynthesis is the biological conversion of one or more carbon-containing molecules (usually carbon dioxide or methane) and nutrients into organic matter using the oxidation of inorganic compounds (e.g., hydrogen gas, hydrog ...
forms the foundation of the food chain. Caves that are close to the surface, such as
lava tube
A lava tube, more rarely called a pyroduct, is a 'roofed conduit through which molten lava travels away from its vent'. If lava in the tube drains out, it will leave an empty cave. Lava tubes are common in low-viscosity volcanic systems. La ...
s, often have tree roots hanging from the cave roof, which provide nutrients for sap-feeding insects. Other important food sources in underground habitats are animals being
decomposed and bat
guano
Guano (Spanish from ) is the accumulated excrement of seabirds or bats. Guano is a highly effective fertiliser due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. Guano was also, to a le ...
,
that creates large
invertebrate
Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
communities
A community is a Level of analysis, social unit (a group of people) with a shared socially-significant characteristic, such as place (geography), place, set of Norm (social), norms, culture, religion, values, Convention (norm), customs, or Ide ...
in such caves.
Ecological classification
Cave dwelling animals show different levels of
adaptation
In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
s 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 natural science of the relationships among living organisms and their Natural environment, environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosystem, and biosphere lev ...
:
*
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);
*
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. Among many scientists, trogloxenes and ...
s: species only occurring sporadically in a hypogean habitat and unable to establish a subterranean population.
Regarding
stygofauna
Stygofauna are any fauna that live in groundwater systems or aquifers, such as caves, fissures and vugs. Stygofauna and troglofauna are the two types of subterranean fauna (based on life-history). Both are associated with subterranean environmen ...
, the corresponding words stygobionts (or stygobites), stygophiles and stygoxenes are used.
Biology
The cave beetle '' '' (family 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
adaptation
In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
s, both
morphological and Physiology">physiological
Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
. 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
''Arachnocampa'' is a genus of nine fungus gnat species which have a bioluminescent larval stage, akin to the larval stage of glowworm beetles. The species of ''Arachnocampa'' are endemic to Australia and New Zealand, dwelling in caves and grotto ...
'' which they detect visually.
Other adaptations include the development and elongation of
antennal 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 unique form of matter with constant chemical composition and characteristic properties. Chemical substances may take the form of a single element or chemical compounds. If two or more chemical substances can be combin ...
,
tactile and humidity
receptors (such as Hamann's organ in the cave
beetle
Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
''Leptodirus'' ''hochenwartii'').
Physiological adaptations include slow
metabolism
Metabolism (, from ''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 cellular processes; the co ...
and reduced
energy
Energy () is the physical quantity, quantitative physical property, property that is transferred to a physical body, body or to a physical system, recognizable in the performance of Work (thermodynamics), work and in the form of heat and l ...
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
ectotherm
An ectotherm (), more commonly referred to as a "cold-blooded animal", is an animal in which internal physiological sources of heat, such as blood, are of relatively small or of quite negligible importance in controlling body temperature.Dav ...
y. As a consequence, cave dwelling animals can resist without eating for long time, live more than comparable epigean species,
reproduce
Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: asexual and sexual.
In asexual reprod ...
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 ...
s.
Evolution and ecology
Subterranean fauna have evolved in isolation.
Stratigraphic
Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks.
Stratigraphy has three related subfields: lithost ...
barriers, such as rock walls and layers, and
fluvial
A river is a natural stream of fresh water that flows on land or inside caves towards another body of water at a lower elevation, such as an ocean, lake, or another river. A river may run dry before reaching the end of its course if it ru ...
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 chemical composition, composition, structure, physical property, physical properties, history, ecology, and the processes by which they form (speleogenes ...
*
Subterranean river
A subterranean river (also known as an underground river) is a river or watercourse that runs wholly or partly beneath the ground, one where the riverbed does not represent the surface of the Earth. It is distinct from an aquifer, which may flow ...
*
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. Among many scientists, trogloxenes and ...
References
{{reflist, 24em
Cave animals
Subterranea (geography)
Animals by adaptation
Animals by location
Ecology