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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 hypogean fish.[1][2]

There are more than 200 scientifically described species of obligate cavefish found on all continents, except Antarctica.[3][4] Although widespread as a group, many cavefish species have very small ranges and are seriously threatened.[5][6] Cavefish are members of a wide range of families and do not form a monophyletic group.[7] Typical adaptations found in cavefish are reduced eyes and pigmentation.[1][2]


Many aboveground fish may enter caves on occasion, but obligate cavefish (fish that require underground habitats) are extremophiles with a number of unusual adaptations known as troglomorphism. In some species, notably the Mexican tetra, shortfin molly, Oman garra, Indoreonectes evezardi and a few catfish, both “normal” aboveground and cavefish forms exist.[10][11][12][13]

Many adaptions seen in cavefish are aimed at surviving in a habitat with little food.[1] Living in darkness, pigmentation and eyes are useless, or an actual disadvantage because of their energy requirements, and therefore typically reduced in cavefish.[14][15][16] Other examples of adaptations are larger fins for more energy-efficient swimming, and a loss of scales and swim bladder.[17][18] The loss can be complete or only partial, for example resulting in small or incomplete (but still existing) eyes, and eyes can be present in the earliest life stages but degenerated by the adult stage.[19] In some cases, “blind” cavefish may still be able to see: Juvenile Mexican tetras of the cave form are able to sense light via certain cells in the pineal gland (pineal eye),[20] and Congo blind barbs are photophobic, despite only having retinas and optical nerves that are rudimentary and located deep inside the head, and completely lacking a lens.[21] In the most extreme cases, the lack of light has changed the circadian rhythm (24-hour internal body clock) of the cavefish. In the Mexican tetra of the cave form and in Phreatichthys andruzzii the circadian rhythm lasts 30 hours and 47 hours, respectively.[22][23] This may help them to save energy.[22] Without sight, other senses are used and these may be enhanced. Examples include the lateral line for sensing vibrations,[24][25][26] mouth suction to sense nearby obstacles (comparable to echolocation),[27] and chemoreception (via smell and taste buds).[28][29] Although there are cavefish in groups known to have electroreception (catfish and South American knifefish), there is no published evidence that this is enhanced in the cave-dwellers.[30] The level of specialized adaptations in a cavefish is generally considered to be directly correlated to the amount of time it has been restricted to the underground habitat: Species that recently arrived show few adaptations and species with the largest number of adaptations are likely the ones that have been restricted to the habitat for the longest time.[31]

Some fish species that live buried in the bottom of aboveground waters, live deep in the sea or live in deep rivers have adaptations similar to cavefish, including reduced eyes and pigmentation.[32][33][34]

Cavefish are quite small with most species being between 2 and 13 cm (0.8–5.1 in) in standard length and about a dozen species reaching 20–23 cm (8–9 in). Only three species grow larger; two slender Ophisternon swamp eels at up to 32–36 cm (13–14 in) in standard length and a much more robust undescribed species of mahseer at 43 cm (17 in).[36][37] The very limited food resources in the habitat likely prevents larger cavefish species from existing and also means that cavefish in general are opportunistic feeders, taking whatever is available.[15][31] In their habitat, cavefish are often the top predators, feeding on smaller cave-living invertebrates, or are detritivores without enemies.[18] Cavefish typically have low metabolic rates and may be able to survive long periods of starvation. A captive Phreatobius cisternarum did not feed for a year, but remained in good condition.[38] The cave form of the Mexican tetra can build up unusually large fat reserves by “binge eating” in periods where food is available, which then (together with its low metabolic rate) allows it to survive without food for months, much longer than the aboveground form of the species.[39]

In the dark habitat, certain types of displays are reduced in cavefish,[17] but in other cases they have become stronger, shifting from displays that are aimed at being seen to displays aimed at being felt via water movement. For example, during the courtship of the cave form of the Mexican tetra the pair produce turbulence through exaggerated gill and mouth movements, allowing them to detect each other.[16] In general, cavefish are slow growers and slow breeders.[2] Breeding behaviors among cavefish vary extensively, and there are both species that are egg-layers and ovoviviparous species that give birth to live young.[16] Uniquely among fish, the genus Amblyopsis brood their eggs in the gill chambers (somewhat like mouthbrooders).[40]


Although many cavefish species are restricted to underground lakes, pools or rivers in actual caves, some are found in aquifers and may only be detected by humans when artificial wells are dug into this layer.[38][41] Most live in areas with low (essentially static) or moderate water current,[1][31] but there are also species in places with very strong current, such as the waterfall climbing cavefish.[42] Underground waters are often very stable environments with limited variations in temperature (typically near the annual average of the surrounding region), nutrient levels and other factors.[1][43] Organic compounds generally only occur in low levels and rely on outside sources, such as contained in water that enters the underground habitat from outside, aboveground animals that find their way into caves (deliberately or by mistake) and guano from bats that roost in caves.[1][43][44] Cavefish are primarily restricted to freshwater.[1] A few species, notably the cave-dwelling viviparous brotulas, Luciogobius gobies, Milyeringa sleeper gobies and the blind cave eel, live in anchialine caves and several of these tolerate various salinities.[1][45][46][47][48]

Range and diversity[edit]

The more than 200 scientifically described obligate cavefish species are found in most continents, but there are strong geographic patterns and the species richness varies.[3] The vast majority of species are found in the tropics or subtropics.[49] Cavefish are strongly linked to regions with karst, which commonly result in underground sinkholes and subterranean rivers.[1][7]

With more than 120 described species, by far the greatest diversity is in Asia, followed by more than 30 species in South America and about 30 species in North America.[3][7] In contrast, only 9 species are known from Africa, 5 from Oceania,[7] and 1 from Europe.[4][50] On a country level, China has the greatest diversity with more than 80 species, followed by Brazil with more than 20 species. India, Mexico, Thailand and the United States of America each have 9–14 species.[1][3][51] No other country has more than 5 cavefish species.[7][52][53]

Being underground, many places where cavefish may live have not been thoroughly surveyed. New cavefish species are described with some regularity and undescribed species are known.[5][7] As a consequence, the number of known cavefish species has risen rapidly in recent decades. In the early 1990s only about 50 species were known, in 2010 about 170 species were known,[55] and by 2015 this had surpassed 200 species.[3] It has been estimated that the final number might be around 250 obligate cavefish species.[56] For example, the first cavefish in Europe, a Barbatula stone loach, was only discovered in 2015 in Southern Germany,[4][50] and the largest known cavefish, Neolissochilus pnar (originally thought to be a form of the golden mahseer), was only definitely confirmed in 2019, despite being quite numerous in the cave where it occurs in Meghalaya, India.[36][37][57] Conversely, their unusual appearance means that some cavefish already attracted attention in ancient times. The oldest known description of an obligate cavefish, involving Sinocyclocheilus hyalinus, is almost 500 years old.[49]

Obligate cavefish are known from a wide range of families: Characidae (characids), Balitoridae (hillstream loaches), Cobitidae (true loaches), Cyprinidae (carps and allies), Nemacheilidae (stone loaches), Amblycipitidae (torrent catfishes), Astroblepidae (naked sucker-mouth catfishes), Callichthyidae (armored catfishes), Clariidae (airbreathing catfishes), Heptapteridae (heptapterid catfishes), Ictaluridae (ictalurid catfishes), Kryptoglanidae (kryptoglanid catfish), Loricariidae (loricariid catfishes), Phreatobiidae (phreatobiid catfishes), Trichomycteridae (pencil catfishes), Sternopygidae (glass knifefishes), Amblyopsidae (U.S. cavefishes), Bythitidae (brotulas), Poeciliidae (live-bearers), Synbranchidae (swamp eels), Cottidae (true sculpins), Butidae (butid gobies), Eleotridae (sleeper gobies), Milyeringidae (blind cave gobies), Gobiidae (gobies) and Channidae (snakeheads).[1][7][58][59][60] Many of these families are only very distantly related and do not form a monophyletic group, showing that adaptations to a life in caves has happened numerous times among fish. As such, their similar adaptions are examples of convergent evolution and the descriptive term “cavefish” is an example of folk taxonomy rather than scientific taxonomy.[7] Strictly speaking some Cyprinodontidae (pupfish) are also known from sinkhole caves, famously including the Devils Hole pupfish, but these lack the adaptations (e.g., reduced eyes and pigmentation) typically associated with cavefish.[1] Additionally, species from a few families such as Chaudhuriidae (earthworm eels), Glanapteryginae and Sarcoglanidinae live buried in the bottom of aboveground waters, and can show adaptions similar to traditional underground-living (troglobitic) fish.[38][32][61][62] It has been argued that such species should be recognized as a part of the group of troglobitic fish.[3]


As of 2019[update], the following underground-living fish species with various levels of troglomorphism (ranging from complete loss of eyes and pigment, to only a partial reduction of one of these) are known.[1][3][51][63] Phreatobius sanguijuela and Prietella phreatophila, the only species with underground populations in more than one country,[64][65] are listed twice. Excluded from the table are species that live buried in the bottom of aboveground waters (even if they have troglomorphic-like features) and undescribed species.

Family Species Country Year of description Notes
Characidae Astyanax aeneus Mexico 1860 Species includes both aboveground and belowground forms (aboveground also in Central America). Sometimes considered a part of Astyanax mexicanus[66][67][68]
Characidae Astyanax mexicanus (blind cave tetra) Mexico 1853 Species includes both aboveground and belowground forms (aboveground also in United States). Cave form sometimes considered a separate species, A. jordani[68]
Characidae Stygichthys typhlops (Brazilian blind characid) Brazil 1965
Cyprinidae Anchicyclocheilus halfibindus China 1992 Sometimes considered a species in the genus Sinocyclocheilus,[63] or a synonym of Sinocyclocheilus microphthalmus[69]
Cyprinidae Barbodes microps Indonesia 1868 Formerly placed in Barbus or Puntius instead. Aboveground populations have also been assigned to this species,[70] but its taxonomy is unresolved and a review has suggested that at least some of the underground populations might belong to Puntius binotatus or an undescribed species instead.[71][72]
Cyprinidae Barbopsis devecchi (Somalian blind barb) Somalia 1926
Cyprinidae Caecobarbus geertsii (Congo blind barb) DR Congo 1921
Cyprinidae Caecocypris basimi (Haditha cavefish) Iraq 1980
Cyprinidae Garra barreimiae (Omani blind cavefish) Oman 1956 Species includes both aboveground and belowground forms (aboveground also in the United Arab Emirates). A population in the United Arab Emirates has been reported to be underground,[51] but this is incorrect[3]
Cyprinidae Garra dunsirei (Tawi Atair garra) Oman 1987
Cyprinidae Garra lorestanensis Iran 2016
Cyprinidae Garra tashanensis Iran 2016
Cyprinidae Garra typhlops (Iran cave barb) Iran 1944 Formerly in its own genus Iranocypris[73]
Cyprinidae Garra widdowsoni (Iraq blind barb) Iraq 1955 Formerly in its own genus Typhlogarra, but genetics show that it belongs in Garra[74][75]
Cyprinidae Longanalus macrochirous China 2006
Cyprinidae Neolissochilus pnar India 2023 Originally tentatively identified as a troglobitic form of the golden mahseer.[57]
Cyprinidae Neolissochilus subterraneus Thailand 2003
Cyprinidae Phreatichthys andruzzii Somalia 1924
Cyprinidae Poropuntius speleops Thailand 1991
Cyprinidae Sinocyclocheilus albeoguttatus China 1998
Cyprinidae Sinocyclocheilus altishoulderus China 1992
Cyprinidae Sinocyclocheilus aluensis China 2005
Cyprinidae Sinocyclocheilus anatirostris (duck-billed golden-line fish) China 1986
Cyprinidae Sinocyclocheilus angularis (gold-colored angle fish) China 1990
Cyprinidae Sinocyclocheilus anophthalmus (eyeless golden-line fish) China 1988
Cyprinidae Sinocyclocheilus anshuiensis China 2013
Cyprinidae Sinocyclocheilus aquihornes China 2007
Cyprinidae Sinocyclocheilus biangularis China 1996
Cyprinidae Sinocyclocheilus bicornutus China 1997
Cyprinidae Sinocyclocheilus brevibarbatus China 2009
Cyprinidae Sinocyclocheilus broadihornes China 2007
Cyprinidae Sinocyclocheilus cyphotergous China 1988
Cyprinidae Sinocyclocheilus flexuosdorsalis China 2012
Cyprinidae Sinocyclocheilus furcodorsalis (crossed-fork back golden-line fish) China 1997
Cyprinidae Sinocyclocheilus guanyangensis China 2016
Cyprinidae Sinocyclocheilus huanjiangensis China 2010
Cyprinidae Sinocyclocheilus hugeibarbus China 2003
Cyprinidae Sinocyclocheilus hyalinus (hyaline fish) China 1993
Cyprinidae Sinocyclocheilus jinxiensis China 2012 Proposed moved to monotypic genus Pseudosinocyclocheilus in 2016[76]
Cyprinidae Sinocyclocheilus jiuxuensis China 2003
Cyprinidae Sinocyclocheilus lingyunensis China 2000
Cyprinidae Sinocyclocheilus longibarbatus China 1989
Cyprinidae Sinocyclocheilus longifinus China 1996
Cyprinidae Sinocyclocheilus luolouensis China 2013
Cyprinidae Sinocyclocheilus luopingensis China 2002
Cyprinidae Sinocyclocheilus macrophthalmus China 2001
Cyprinidae Sinocyclocheilus macroscalus China 2000
Cyprinidae Sinocyclocheilus maculatus China 2000
Cyprinidae Sinocyclocheilus maitianheensis China 1992
Cyprinidae Sinocyclocheilus malacopterus China 1985
Cyprinidae Sinocyclocheilus mashanensis China 2010
Cyprinidae Sinocyclocheilus microphthalmus (small eye golden-line fish) China 1989
Cyprinidae Sinocyclocheilus multipunctatus China 1931
Cyprinidae Sinocyclocheilus oxycephalus China 1985
Cyprinidae Sinocyclocheilus purpureus China 1985
Cyprinidae Sinocyclocheilus qiubeiensis China 2002
Cyprinidae Sinocyclocheilus rhinocerous China 1994
Cyprinidae Sinocyclocheilus robustus China 1988
Cyprinidae Sinocyclocheilus tianeensis China 2003
Cyprinidae Sinocyclocheilus tianlinensis China 2004
Cyprinidae Sinocyclocheilus tileihornes China 2003
Cyprinidae Sinocyclocheilus xunlensis China 2004
Cyprinidae Sinocyclocheilus yishanensis China 1992
Cyprinidae Speolabeo hokhanhi Vietnam 2018
Cyprinidae Speolabeo musaei Laos 2011 Formerly in genus Bangana[77]
Cyprinidae Troglocyclocheilus khammouanensis Laos 1999
Cyprinidae Typhlobarbus nudiventris China 1982
Balitoridae Cryptotora thamicola (waterfall climbing cavefish) Thailand 1988
Cobitidae Bibarba parvoculus China 2015
Cobitidae Cobitis damlae Turkey 2014 First described as a species of cavefish based on a single specimen, but a later review suggested that it was found in an area without underground waters and only is an albinistic individual of the aboveground Cobitis fahireae[78]
Cobitidae Pangio bhujia India 2019
Cobitidae Protocobitis anteroventris China 2013
Cobitidae Protocobitis polylepis China 2008
Cobitidae Protocobitis typhlops China 1993
Nemacheilidae Barbatula barbatula (stone loach) Germany 1758 Aboveground populations widespread in Europe. Belowground population only discovered in 2015 and tentatively included in this species based on genetic evidence. Only known cavefish in Europe[4]
Nemacheilidae Claea dabryi China 1874 Traditionally in genus Schistura or Triplophysa.[79][80] Species includes both aboveground and belowground populations; the latter sometimes recognized as a separate subspecies microphthalmus.[63]
Nemacheilidae Draconectes narinosus Vietnam 2012
Nemacheilidae Eidinemacheilus proudlovei Iraq 2016
Nemacheilidae Eidinemacheilus smithi (Zagroz blind loach) Iran 1976 Formerly in genus Noemacheilus or Paracobitis[81]
Nemacheilidae Heminoemacheilus hyalinus China 1996
Nemacheilidae Indoreonectes evezardi India 1872 Species includes both aboveground and belowground forms[82]
Nemacheilidae Nemacheilus troglocataractus (blind cave loach) Thailand 1989
Nemacheilidae Oreonectes acridorsalis China 2013
Nemacheilidae Oreonectes anophthalmus China 1981
Nemacheilidae Oreonectes barbatus China 2013
Nemacheilidae Oreonectes daqikongensis China 2016
Nemacheilidae Oreonectes donglanensis China 2013
Nemacheilidae Oreonectes duanensis China 2013
Nemacheilidae Oreonectes elongatus China 2012
Nemacheilidae Oreonectes furcocaudalis China 1987
Nemacheilidae Oreonectes guananensis China 2011
Nemacheilidae Oreonectes luochengensis China 2011
Nemacheilidae Oreonectes macrolepis China 2009
Nemacheilidae Oreonectes microphthalmus China 2008
Nemacheilidae Oreonectes shuilongensis China 2016
Nemacheilidae Oreonectes translucens China 2006
Nemacheilidae Schistura deansmarti Thailand 2003
Nemacheilidae Schistura jarutanini Thailand 1990
Nemacheilidae Schistura kaysonei Laos 2002
Nemacheilidae Schistura larketensis India 2017
Nemacheilidae Schistura lingyunensis China 1997 Sometimes in genus Triplophysa[79]
Nemacheilidae Schistura mobbsi Vietnam 2012
Nemacheilidae Schistura oedipus Thailand 1988
Nemacheilidae Schistura papulifera India 2007
Nemacheilidae Schistura sijuensis India 1987
Nemacheilidae Schistura spekuli Vietnam 2004
Nemacheilidae Schistura spiesi Thailand 2003
Nemacheilidae Speonectes tiomanensis Malaysia 1990 Formerly in genus Sundoreonectes[79]
Nemacheilidae Triplophysa aluensis China 2000
Nemacheilidae Triplophysa dongganensis China 2013
Nemacheilidae Triplophysa fengshanensis China 2013
Nemacheilidae Triplophysa gejiuensis China 1979
Nemacheilidae Triplophysa huanjiangensis China 2011
Nemacheilidae Triplophysa jiarongensis China 2012
Nemacheilidae Triplophysa langpingensis China 2013
Nemacheilidae Triplophysa lihuensis China 2012
Nemacheilidae Triplophysa longibarbata China 1998 Includes Paracobitis maolanensis and P. posterodorsalus as synonyms,[79] which may be valid species[63]
Nemacheilidae Triplophysa luochengensis China 2017
Nemacheilidae Triplophysa macrocephala China 2011
Nemacheilidae Triplophysa qiubeiensis China 2008
Nemacheilidae Triplophysa rosa China 2005
Nemacheilidae Triplophysa shilinensis China 1992
Nemacheilidae Triplophysa tianeensis China 2004
Nemacheilidae Triplophysa xiangshuingensis China 2004
Nemacheilidae Triplophysa xiangxiensis China 1986
Nemacheilidae Triplophysa yunnanensis China 1990
Nemacheilidae Troglocobitis starostini (Starostin’s loach) Turkmenistan 1983
Amblycipitidae Xiurenbagrus dorsalis China 2014
Astroblepidae Astroblepus pholeter Ecuador 1962
Astroblepidae Astroblepus riberae Peru 1994
Callichthyidae Aspidoras mephisto Brazil 2017 Formerly included in aboveground species A. albater[83]
Clariidae Clarias cavernicola (golden cave catfish) Angola 1936
Clariidae Horaglanis abdulkalami India 2012
Clariidae Horaglanis alikunhii India 2004
Clariidae Horaglanis krishnai (Indian blind catfish) India 1950
Clariidae Uegitglanis zammaranoi Somalia 1923
Heptapteridae Pimelodella kronei Brazil 1907
Heptapteridae Pimelodella spelaea Brazil 2004
Heptapteridae Rhamdia enfurnada Brazil 2005
Heptapteridae Rhamdia guasarensis Venezuela 2004
Heptapteridae Rhamdia laluchensis (La Lucha blind catfish) Mexico 2003
Heptapteridae Rhamdia laticauda typhla Belize 1982 Other subspecies found in aboveground habitats in Mexico and Central America[1][84]
Heptapteridae Rhamdia macuspanensis (Olmec blind catfish) Mexico 1998
Heptapteridae Rhamdia quelen urichi Trinidad 1926 Other subspecies found widely in aboveground habitats in South and Central America[85]
Heptapteridae Rhamdia reddelli (blind whiskered catfish) Mexico 1984
Heptapteridae Rhamdia zongolicensis (Zongolica catfish) Mexico 1993
Heptapteridae Rhamdiopsis krugi Brazil 2010
Ictaluridae Prietella lundbergi (phantom blindcat) Mexico 1995
Ictaluridae Prietella phreatophila (Mexican blindcat) Mexico 1954 Listed twice (once for each country)
Ictaluridae Prietella phreatophila (Mexican blindcat) United States 1954 Listed twice (once for each country)
Ictaluridae Satan eurystomus (widemouth blindcat) United States 1947
Ictaluridae Trogloglanis pattersoni (toothless blindcat) United States 1919
Kryptoglanidae Kryptoglanis shajii India 2011 Found both underground and aboveground (not known to differ in appearance)[86]
Loricariidae Ancistrus cryptophthalmus Brazil 1987
Loricariidae Ancistrus formoso Brazil 1997
Loricariidae Ancistrus galani Venezuela 1994
Phreatobiidae Phreatobius cisternarum Brazil 1905
Phreatobiidae Phreatobius dracunculus Brazil 2007
Phreatobiidae Phreatobius sanguijuela Bolivia 2007 Listed twice (once for each country)
Phreatobiidae Phreatobius sanguijuela Brazil 2007 Listed twice (once for each country)
Siluridae Pterocryptis buccata (cave sheatfish) Thailand 1998 Species includes both aboveground and belowground forms[13]
Siluridae Pterocryptis cucphuongensis Vietnam 1978
Trichomycteridae Glaphyropoma spinosum Brazil 2008
Trichomycteridae Ituglanis bambui Brazil 2004
Trichomycteridae Ituglanis boticario Brazil 2015
Trichomycteridae Ituglanis epikarsticus Brazil 2004
Trichomycteridae Ituglanis mambai Brazil 2008
Trichomycteridae Ituglanis passensis Brazil 2002
Trichomycteridae Ituglanis ramiroi Brazil 2004
Trichomycteridae Silvinichthys bortayro Argentina 2005
Trichomycteridae Trichomycterus dali Brazil 2011
Trichomycteridae Trichomycterus chaberti Bolivia 1968
Trichomycteridae Trichomycterus itacarambiensis Brazil 1996
Trichomycteridae Trichomycterus rosablanca Colombia 2018
Trichomycteridae Trichomycterus rubbioli Brazil 2012
Trichomycteridae Trichomycterus sandovali Colombia 2006
Trichomycteridae Trichomycterus santanderensis Colombia 2007
Trichomycteridae Trichomycterus sketi Colombia 2010
Trichomycteridae Trichomycterus spelaeus Venezuela 2001
Trichomycteridae Trichomycterus uisae (trepador) Colombia 2008
Sternopygidae Eigenmannia vicentespelaea Brazil 1996
Amblyopsidae Amblyopsis hoosieri (Hoosier cavefish) United States 2014
Amblyopsidae Amblyopsis rosae (Ozark cavefish) United States 1898
Amblyopsidae Amblyopsis spelaea (northern cavefish) United States 1842
Amblyopsidae Forbesichthys agassizii (spring cavefish) United States 1872 Found belowground, but also nearby in aboveground waters during the night[1][87]
Amblyopsidae Speoplatyrhinus poulsoni (Alabama cavefish) United States 1974
Amblyopsidae Typhlichthys subterraneus (southern cavefish) United States 1859 Possibly a species complex and T. eigemanni may be a valid species[88]
Bythitidae Diancistrus typhlops Indonesia 2009
Bythitidae Lucifuga dentata (toothed Cuban cusk-eel) Cuba 1858
Bythitidae Lucifuga lucayana (Lucaya cave brotula) Bahamas 2006
Bythitidae Lucifuga simile Cuba 1981
Bythitidae Lucifuga spelaeotes (New Providence cusk-eel) Bahamas 1970
Bythitidae Lucifuga subterranea (Cuban cusk-eel) Cuba 1858
Bythitidae Lucifuga teresinarum Cuba 1988
Bythitidae Ogilbia galapagosensis (Galapagos cuskeel) Ecuador 1965 Arguably not a true cavefish, as places it inhabits also can be described as lagoon crevices[1]
Bythitidae Typhliasina pearsei (Mexican blind brotula) Mexico 1938
Poeciliidae Poecilia mexicana (cave molly) Mexico 1863 Species includes both aboveground and belowground forms (aboveground also in Central America)[10]
Synbranchidae Rakthamichthys digressus India 2002
Synbranchidae Rakthamichthys indicus India 1961 Originally described as Monopterus indicus by K. C. Eapen, but as this name was already taken by the Bombay swamp eel, it was redescribed as Monopterus eapeni in 1991. When the species was moved to the genus Rakthamichthys, the indicus specific epithet was revived.
Synbranchidae Rakthamichthys roseni India 1998
Synbranchidae Ophisternon candidum (blind cave eel) Australia 1962
Synbranchidae Ophisternon infernale (blind swamp eel) Mexico 1938
Cottidae C. bairdi—cognatus species complex (mottled sculpin/slimy sculpin) United States 1850/1836 Aboveground forms relatively widespread in North America and Siberia, underground form only in Pennsylvania[89]
Cottidae Cottus carolinae (banded sculpin) United States 1861 Aboveground forms relatively widespread in the United States, underground form only in West Virginia[90][91]
Cottidae Cottus specus (grotto sculpin) United States 2013 Formerly included in C. carolinae[91]
Butidae Bostrychus microphthalmus Indonesia 2005 The family Butidae was formerly considered a subfamily of Eleotridae[3]
Butidae Oxyeleotris caeca Papua New Guinea 1996 The family Butidae was formerly considered a subfamily of Eleotridae[3]
Butidae Oxyeleotris colasi Indonesia 2013 Has mistakenly been reported to occur in Papua New Guinea,[3] but it is from Western New Guinea, the Indonesian part of the island.[92] The family Butidae was formerly considered a subfamily of Eleotridae[3]
Eleotridae Caecieleotris morrisi (Oaxaca cave sleeper) Mexico 2016
Milyeringidae Milyeringa brooksi Australia 2010 The family Milyeringidae was formerly considered a subfamily of Eleotridae[93]
Milyeringidae Milyeringa justitia (Barrow cave gudgeon) Australia 2013 The family Milyeringidae was formerly considered a subfamily of Eleotridae[93]
Milyeringidae Milyeringa veritas (blind gudgeon) Australia 1945 The family Milyeringidae was formerly considered a subfamily of Eleotridae[93]
Milyeringidae Typhleotris madagascariensis Madagascar 1933 The family Milyeringidae was formerly considered a subfamily of Eleotridae[94]
Milyeringidae Typhleotris mararybe Madagascar 2012 The family Milyeringidae was formerly considered a subfamily of Eleotridae[94]
Milyeringidae Typhleotris pauliani Madagascar 1959 The family Milyeringidae was formerly considered a subfamily of Eleotridae[94]
Gobiidae Caecogobius cryptophthalmus Philippines 1991
Gobiidae Caecogobius personatus Philippines 2019
Gobiidae Glossogobius ankaranensis Madagascar 1994
Gobiidae Luciogobius albus Japan 1940
Gobiidae Luciogobius pallidus Japan 1940
Aenigmachannidae Aenigmachanna gollum (Gollum snakehead) India 2019 One of two species in a unique fish family closely related to true snakeheads. Displays relatively few troglomorphisms despite living in underground aquifers, and thus could either be a recent arrival to the subterranean ecosystem or possibly a subtroglophile that periodically moves between the underground and surface.[59]
Aenigmachannidae Aenigmachanna mahabali India 2019 One of two species in a unique fish family closely related to true snakeheads. Displays relatively few troglomorphisms despite living in underground aquifers, and thus could either be a recent arrival to the subterranean ecosystem or possibly a subtroglophile that periodically moves between the underground and surface.[60]


Although cavefish as a group are found throughout large parts of the world, many cavefish species have tiny ranges (often restricted to a single cave or cave system) and are seriously threatened. In 1996, more than 50 species were recognized as threatened by the IUCN and many, including several that are rare, have not been assessed at all.[2] For example, the critically endangered Alabama cavefish is only found in the Key Cave and the entire population has been estimated at less than 100 individuals,[95] while the critically endangered golden cave catfish only is found in the Aigamas cave in Namibia and has an estimated population of less than 400 individuals.[96] The Haditha cavefish from Iraq and the Oaxaca cave sleeper from Mexico may already be extinct, as recent surveys have failed to find them.[97][98] In some other cases, such as the Brazilian blind characid which went unrecorded by ichthyologists from 1962 to 2004, the apparent “rarity” was likely because of a lack of surveys in its range and habitat, as locals considered it relatively common until the early 1990s (more recently, this species appears to truly have declined significantly).[41] Living in very stable environments, cavefish are likely more vulnerable to changes in the water (for example, temperature or oxygen) than fish of aboveground habitats which naturally experience greater variations.[43] The main threats to cavefish are typically changes in the water level (mainly through water extraction or drought), habitat degradation and pollution, but in some cases introduced species and collection for the aquarium trade also present a threat.[5][6] Cavefish often show little fear of humans and can sometimes be caught with the bare hands.[18] Most cavefish lack natural predators, although larger cavefish may feed on smaller individuals,[18] and cave-living crayfish, crabs, giant water bugs and spiders have been recorded feeding on a few species of cavefish.[99][100][101][102]

Caves in some parts of the world have been protected, which can safeguard the cavefish.[54] In a few cases such as the Omani blind cavefish (Oman garra), zoos have initiated breeding programs as a safeguard.[12] In contrast to the rarer species, the cave form of the Mexican tetra is easily bred in captivity and widely available to aquarists.[68][103] This is the most studied cavefish species and likely also the most studied cave organism overall.[104] As of 2006, only six other cavefish species have been bred in captivity, typically by scientists.[56]

See also[edit]


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