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Cavefish

April 07, 2023

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]

Phreatichthys andruzzii showing the pale colour and lack of eyes typical of cavefish. The large red spot on the head is the blood-filled gills, visible through the semi-transparent gill cover

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]

Adaptations

 
As typical of cavefish, Typhleotris madagascariensis is an opportunistic feeder on various invertebrates[8][9]

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]

 
The waterfall climbing cavefish has several adaptions that allow it to climb and "walk" in a tetrapod-like fashion[35]

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]

Habitat

 
The Mexican blind brotula and other cave-dwelling brotulas are among the few species that live in anchialine habitats

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

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]

 
The Hoosier cavefish from Indiana in the United States was only described in 2014[54]

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]

Species

As of 2019, 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.

Conservation

 
The cave form of the Mexican tetra is easily bred in captivity and the only cavefish widely available to aquarists

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

References

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April 07, 2023
cavefish, cave, fish, generic, term, fresh, brackish, water, fish, adapted, life, caves, other, underground, habitats, related, terms, subterranean, fish, troglomorphic, fish, troglobitic, fish, stygobitic, fish, phreatic, fish, hypogean, fish, phreatichthys, . 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 Phreatichthys andruzzii showing the pale colour and lack of eyes typical of cavefish The large red spot on the head is the blood filled gills visible through the semi transparent gill cover 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 Contents 1 Adaptations 2 Habitat 3 Range and diversity 3 1 Species 4 Conservation 5 See also 6 ReferencesAdaptations Edit As typical of cavefish Typhleotris madagascariensis is an opportunistic feeder on various invertebrates 8 9 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 The waterfall climbing cavefish has several adaptions that allow it to climb and walk in a tetrapod like fashion 35 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 Habitat Edit The Mexican blind brotula and other cave dwelling brotulas are among the few species that live in anchialine habitats 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 EditThe 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 The Hoosier cavefish from Indiana in the United States was only described in 2014 54 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 Species Edit 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 NotesCharacidae 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 1965Cyprinidae 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 1926Cyprinidae Caecobarbus geertsii Congo blind barb DR Congo 1921Cyprinidae Caecocypris basimi Haditha cavefish Iraq 1980Cyprinidae 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 1987Cyprinidae Garra lorestanensis Iran 2016Cyprinidae Garra tashanensis Iran 2016Cyprinidae 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 2006Cyprinidae Neolissochilus pnar India 2023 Originally tentatively identified as a troglobitic form of the golden mahseer 57 Cyprinidae Neolissochilus subterraneus Thailand 2003Cyprinidae Phreatichthys andruzzii Somalia 1924Cyprinidae Poropuntius speleops Thailand 1991Cyprinidae Sinocyclocheilus albeoguttatus China 1998Cyprinidae Sinocyclocheilus altishoulderus China 1992Cyprinidae Sinocyclocheilus aluensis China 2005Cyprinidae Sinocyclocheilus anatirostris duck billed golden line fish China 1986Cyprinidae Sinocyclocheilus angularis gold colored angle fish China 1990Cyprinidae Sinocyclocheilus anophthalmus eyeless golden line fish China 1988Cyprinidae Sinocyclocheilus anshuiensis China 2013Cyprinidae Sinocyclocheilus aquihornes China 2007Cyprinidae Sinocyclocheilus biangularis China 1996Cyprinidae Sinocyclocheilus bicornutus China 1997Cyprinidae Sinocyclocheilus brevibarbatus China 2009Cyprinidae Sinocyclocheilus broadihornes China 2007Cyprinidae Sinocyclocheilus cyphotergous China 1988Cyprinidae Sinocyclocheilus flexuosdorsalis China 2012Cyprinidae Sinocyclocheilus furcodorsalis crossed fork back golden line fish China 1997Cyprinidae Sinocyclocheilus guanyangensis China 2016Cyprinidae Sinocyclocheilus huanjiangensis China 2010Cyprinidae Sinocyclocheilus hugeibarbus China 2003Cyprinidae Sinocyclocheilus hyalinus hyaline fish China 1993Cyprinidae Sinocyclocheilus jinxiensis China 2012 Proposed moved to monotypic genus Pseudosinocyclocheilus in 2016 76 Cyprinidae Sinocyclocheilus jiuxuensis China 2003Cyprinidae Sinocyclocheilus lingyunensis China 2000Cyprinidae Sinocyclocheilus longibarbatus China 1989Cyprinidae Sinocyclocheilus longifinus China 1996Cyprinidae Sinocyclocheilus luolouensis China 2013Cyprinidae Sinocyclocheilus luopingensis China 2002Cyprinidae Sinocyclocheilus macrophthalmus China 2001Cyprinidae Sinocyclocheilus macroscalus China 2000Cyprinidae Sinocyclocheilus maculatus China 2000Cyprinidae Sinocyclocheilus maitianheensis China 1992Cyprinidae Sinocyclocheilus malacopterus China 1985Cyprinidae Sinocyclocheilus mashanensis China 2010Cyprinidae Sinocyclocheilus microphthalmus small eye golden line fish China 1989Cyprinidae Sinocyclocheilus multipunctatus China 1931Cyprinidae Sinocyclocheilus oxycephalus China 1985Cyprinidae Sinocyclocheilus purpureus China 1985Cyprinidae Sinocyclocheilus qiubeiensis China 2002Cyprinidae Sinocyclocheilus rhinocerous China 1994Cyprinidae Sinocyclocheilus robustus China 1988Cyprinidae Sinocyclocheilus tianeensis China 2003Cyprinidae Sinocyclocheilus tianlinensis China 2004Cyprinidae Sinocyclocheilus tileihornes China 2003Cyprinidae Sinocyclocheilus xunlensis China 2004Cyprinidae Sinocyclocheilus yishanensis China 1992Cyprinidae Speolabeo hokhanhi Vietnam 2018Cyprinidae Speolabeo musaei Laos 2011 Formerly in genus Bangana 77 Cyprinidae Troglocyclocheilus khammouanensis Laos 1999Cyprinidae Typhlobarbus nudiventris China 1982Balitoridae Cryptotora thamicola waterfall climbing cavefish Thailand 1988Cobitidae Bibarba parvoculus China 2015Cobitidae 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 2019Cobitidae Protocobitis anteroventris China 2013Cobitidae Protocobitis polylepis China 2008Cobitidae Protocobitis typhlops China 1993Nemacheilidae 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 2012Nemacheilidae Eidinemacheilus proudlovei Iraq 2016Nemacheilidae Eidinemacheilus smithi Zagroz blind loach Iran 1976 Formerly in genus Noemacheilus or Paracobitis 81 Nemacheilidae Heminoemacheilus hyalinus China 1996Nemacheilidae Indoreonectes evezardi India 1872 Species includes both aboveground and belowground forms 82 Nemacheilidae Nemacheilus troglocataractus blind cave loach Thailand 1989Nemacheilidae Oreonectes acridorsalis China 2013Nemacheilidae Oreonectes anophthalmus China 1981Nemacheilidae Oreonectes barbatus China 2013Nemacheilidae Oreonectes daqikongensis China 2016Nemacheilidae Oreonectes donglanensis China 2013Nemacheilidae Oreonectes duanensis China 2013Nemacheilidae Oreonectes elongatus China 2012Nemacheilidae Oreonectes furcocaudalis China 1987Nemacheilidae Oreonectes guananensis China 2011Nemacheilidae Oreonectes luochengensis China 2011Nemacheilidae Oreonectes macrolepis China 2009Nemacheilidae Oreonectes microphthalmus China 2008Nemacheilidae Oreonectes shuilongensis China 2016Nemacheilidae Oreonectes translucens China 2006Nemacheilidae Schistura deansmarti Thailand 2003Nemacheilidae Schistura jarutanini Thailand 1990Nemacheilidae Schistura kaysonei Laos 2002Nemacheilidae Schistura larketensis India 2017Nemacheilidae Schistura lingyunensis China 1997 Sometimes in genus Triplophysa 79 Nemacheilidae Schistura mobbsi Vietnam 2012Nemacheilidae Schistura oedipus Thailand 1988Nemacheilidae Schistura papulifera India 2007Nemacheilidae Schistura sijuensis India 1987Nemacheilidae Schistura spekuli Vietnam 2004Nemacheilidae Schistura spiesi Thailand 2003Nemacheilidae Speonectes tiomanensis Malaysia 1990 Formerly in genus Sundoreonectes 79 Nemacheilidae Triplophysa aluensis China 2000Nemacheilidae Triplophysa dongganensis China 2013Nemacheilidae Triplophysa fengshanensis China 2013Nemacheilidae Triplophysa gejiuensis China 1979Nemacheilidae Triplophysa huanjiangensis China 2011Nemacheilidae Triplophysa jiarongensis China 2012Nemacheilidae Triplophysa langpingensis China 2013Nemacheilidae Triplophysa lihuensis China 2012Nemacheilidae Triplophysa longibarbata China 1998 Includes Paracobitis maolanensis and P posterodorsalus as synonyms 79 which may be valid species 63 Nemacheilidae Triplophysa luochengensis China 2017Nemacheilidae Triplophysa macrocephala China 2011Nemacheilidae Triplophysa qiubeiensis China 2008Nemacheilidae Triplophysa rosa China 2005Nemacheilidae Triplophysa shilinensis China 1992Nemacheilidae Triplophysa tianeensis China 2004Nemacheilidae Triplophysa xiangshuingensis China 2004Nemacheilidae Triplophysa xiangxiensis China 1986Nemacheilidae Triplophysa yunnanensis China 1990Nemacheilidae Troglocobitis starostini Starostin s loach Turkmenistan 1983Amblycipitidae Xiurenbagrus dorsalis China 2014Astroblepidae Astroblepus pholeter Ecuador 1962Astroblepidae Astroblepus riberae Peru 1994Callichthyidae Aspidoras mephisto Brazil 2017 Formerly included in aboveground species A albater 83 Clariidae Clarias cavernicola golden cave catfish Angola 1936Clariidae Horaglanis abdulkalami India 2012Clariidae Horaglanis alikunhii India 2004Clariidae Horaglanis krishnai Indian blind catfish India 1950Clariidae Uegitglanis zammaranoi Somalia 1923Heptapteridae Pimelodella kronei Brazil 1907Heptapteridae Pimelodella spelaea Brazil 2004Heptapteridae Rhamdia enfurnada Brazil 2005Heptapteridae Rhamdia guasarensis Venezuela 2004Heptapteridae Rhamdia laluchensis La Lucha blind catfish Mexico 2003Heptapteridae 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 1998Heptapteridae 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 1984Heptapteridae Rhamdia zongolicensis Zongolica catfish Mexico 1993Heptapteridae Rhamdiopsis krugi Brazil 2010Ictaluridae Prietella lundbergi phantom blindcat Mexico 1995Ictaluridae 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 1947Ictaluridae Trogloglanis pattersoni toothless blindcat United States 1919Kryptoglanidae Kryptoglanis shajii India 2011 Found both underground and aboveground not known to differ in appearance 86 Loricariidae Ancistrus cryptophthalmus Brazil 1987Loricariidae Ancistrus formoso Brazil 1997Loricariidae Ancistrus galani Venezuela 1994Phreatobiidae Phreatobius cisternarum Brazil 1905Phreatobiidae Phreatobius dracunculus Brazil 2007Phreatobiidae 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 1978Trichomycteridae Glaphyropoma spinosum Brazil 2008Trichomycteridae Ituglanis bambui Brazil 2004Trichomycteridae Ituglanis boticario Brazil 2015Trichomycteridae Ituglanis epikarsticus Brazil 2004Trichomycteridae Ituglanis mambai Brazil 2008Trichomycteridae Ituglanis passensis Brazil 2002Trichomycteridae Ituglanis ramiroi Brazil 2004Trichomycteridae Silvinichthys bortayro Argentina 2005Trichomycteridae Trichomycterus dali Brazil 2011Trichomycteridae Trichomycterus chaberti Bolivia 1968Trichomycteridae Trichomycterus itacarambiensis Brazil 1996Trichomycteridae Trichomycterus rosablanca Colombia 2018Trichomycteridae Trichomycterus rubbioli Brazil 2012Trichomycteridae Trichomycterus sandovali Colombia 2006Trichomycteridae Trichomycterus santanderensis Colombia 2007Trichomycteridae Trichomycterus sketi Colombia 2010Trichomycteridae Trichomycterus spelaeus Venezuela 2001Trichomycteridae Trichomycterus uisae trepador Colombia 2008Sternopygidae Eigenmannia vicentespelaea Brazil 1996Amblyopsidae Amblyopsis hoosieri Hoosier cavefish United States 2014Amblyopsidae Amblyopsis rosae Ozark cavefish United States 1898Amblyopsidae Amblyopsis spelaea northern cavefish United States 1842Amblyopsidae 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 1974Amblyopsidae Typhlichthys subterraneus southern cavefish United States 1859 Possibly a species complex and T eigemanni may be a valid species 88 Bythitidae Diancistrus typhlops Indonesia 2009Bythitidae Lucifuga dentata toothed Cuban cusk eel Cuba 1858Bythitidae Lucifuga lucayana Lucaya cave brotula Bahamas 2006Bythitidae Lucifuga simile Cuba 1981Bythitidae Lucifuga spelaeotes New Providence cusk eel Bahamas 1970Bythitidae Lucifuga subterranea Cuban cusk eel Cuba 1858Bythitidae Lucifuga teresinarum Cuba 1988Bythitidae 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 1938Poeciliidae Poecilia mexicana cave molly Mexico 1863 Species includes both aboveground and belowground forms aboveground also in Central America 10 Synbranchidae Rakthamichthys digressus India 2002Synbranchidae 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 1998Synbranchidae Ophisternon candidum blind cave eel Australia 1962Synbranchidae Ophisternon infernale blind swamp eel Mexico 1938Cottidae 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 2016Milyeringidae 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 1991Gobiidae Caecogobius personatus Philippines 2019Gobiidae Glossogobius ankaranensis Madagascar 1994Gobiidae Luciogobius albus Japan 1940Gobiidae Luciogobius pallidus Japan 1940Aenigmachannidae 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 Conservation Edit The cave form of the Mexican tetra is easily bred in captivity and the only cavefish widely available to aquarists 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 EditCave salamanderReferences Edit a b c d e f g h i j k l m n o p Romero Aldemaro editor 2001 The Biology of Hypogean Fishes Developments in Environmental Biology of Fishes ISBN 978 1402000768 a b c d Helfman G S 2007 Fish Conservation A Guide to Understanding and Restoring Global Aquatic Biodiversity and Fishery Resources pp 41 42 Island Press ISBN 978 1 55963 595 0 a b c d e f g h i j k l Proudlove G R 2015 Checklist of troglobitic subterranean fishes of the world to February 2015 cave registry org uk Retrieved 14 May 2017 a b c d Behrmann Godel J A W Nolte J Kreiselmaier R Berka J Freyhof 2017 The first European cave fish Current Biology 27 7 R257 R258 doi 10 1016 j cub 2017 02 048 PMID 28376329 a b c Fenolio D B Zhao Y Niemiller M L and Stout J 2013 In situ observations of seven enigmatic cave loaches and one cave barbel from Guangxi China with notes on conservation status Speleobiology Notes 5 19 33 a b Proudlove G S 2001 The conservation of hypogean fishes Environmental Biology of Fishes 62 201 213 a b c d e f g h Riesch R Tobler M and Plath M 2015 Extremophile Fishes Ecology Evolution and Physiology of Teleosts in Extreme Environments ISBN 978 3319133614 Froese Rainer Pauly Daniel eds 2017 Typhleotris madagascariensis in FishBase April 2017 version Rasoloariniaina Ganzhorn Riemann amp Raminosoa 2016 Water quality and biotic interaction of two cavefish species Typhleotris madagascariensis Petit 1933 and Typhleotris mararybe Sparks amp Chakrabarty 2012 in the Mahafaly Plateau groundwater system Madagascar Subterranean Biology 18 1 16 doi 10 3897 subtbiol 18 8321 a b Plath M and Tobler M 2007 Sex recognition in surface and cave dwelling Atlantic molly females Poecilia mexicana Poeciliidae Teleostei influence of visual and non visual cues acta ethol 10 81 88 Gross J B 2012 The complex origin of Astyanax cavefish BMC Evolutionary Biology 12 105 a b Harrison I J 2015 Garra barreimiae IUCN Red List of Threatened Species 2015 e T8916A3147989 doi 10 2305 IUCN UK 2015 2 RLTS T8916A3147989 en Retrieved 23 December 2017 a b Ng H H and Kottelat M 1998 Pterocryptis buccata a new species of catfish from western Thailand Teleostei Siluridae with epigean and hypogean populations Ichthyological Research 45 4 393 399 Rantin B Bichuette M E 2013 Phototactic behaviour of subterranean Copionodontinae Pinna 1992 catfishes Siluriformes Trichomycteridae from Chapada Diamantina central Bahia northeastern Brazil International Journal of Speleology 41 1 57 63 doi 10 5038 1827 806X 42 1 7 a b Owen J 11 September 2015 How This Cave Dwelling Fish Lost Its Eyes to Evolution National Geographic Retrieved 14 May 2017 a b c Burton M et al 2002 International Wildlife Encyclopedia volume 3 Bro Che 3rd ed p 410 ISBN 978 2 9527084 0 1 a b Romero S and Green S M 2005 The end of regressive evolution examining and interpreting the evidence from cave fishes Journal of Fish Biology 67 1 3 32 a b c d Parzefall J and Trajano E 2010 Behavioral Patterns in Subterranean Fishes In Trajano E Bichuette M E and Kapoor B G eds Biology of Subterranean Fishes ISBN 978 1578086702 Secutti S amp E Trajano 2009 Reproductive behavior development and eye regression in the cave armored catfish Ancistrus cryptophthalmus Reis 1987 Siluriformes Loricariidae breed in laboratory Neotropical Ichthyology 7 3 479 490 doi 10 1590 S1679 62252009000300016 Choi C Q 28 January 2008 Blind Fish Still Able to See LiveScience Retrieved 28 February 2016 Vreven E A Kimbembi ma Ibaka amp S Wamuini Lunkayilakio 2011 The Congo blind barb Mbanza Ngungu s albino cave fish In Darwall Smith Allen Holland Harrison amp Brooks eds The diversity of life in African freshwaters Underwater under threat IUCN pp 74 75 ISBN 978 2 8317 1345 8 a b Palermo E 24 September 2014 Blind Cavefish Froze Its Internal Clock to Save Energy LiveScience Retrieved 28 February 2016 Battison L 10 September 2011 Fish living in dark caves still feel the rhythm of life BBC News Retrieved 28 February 2016 Burt de Perera T 2004 Spatial parameters encoded in the spatial map of the blind Mexican cave fish Astyanax fasciatus Anim Behav 68 291 295 Weber A 1995 The lateral line system of epigean and cave dwelling catfishes of the genus Rhamdia Pimelodidae Teleostei in Mexico Mem Biospeol 22 215 225 Yoshizawa Masato Goricki Spela Soares Daphne Jeffery William R September 2010 Evolution of a Behavioral Shift Mediated by Superficial Neuromasts Helps Cavefish Find Food in Darkness Current Biology 20 18 1631 1636 doi 10 1016 j cub 2010 07 017 PMC 2946428 PMID 20705469 Poppick L 2 April 2014 Mouth Vision Blind Fish Suctions Water to Navigate LiveScience Retrieved 28 February 2016 Bibliowicz J Alie A Espinasa L Yoshizawa M Blin M Hinaux H Legendre L Pere S and Retaux S 2013 Differences in chemosensory response between eyed and eyeless Astyanax mexicanus of the Rio Subterraneo cave EvoDevo 25 Kasumyan A O amp E A Marusov 2015 Chemoorientation in the feeding behavior of the blind Mexican cavefish Astyanax fasciatus Characidae Teleostei Russian Journal of Ecology 46 6 559 563 doi 10 1134 s1067413615060053 S2CID 17283377 Soares D M L Niemiller 2013 Sensory Adaptations of Fishes to Subterranean Environments BioScience 63 4 274 283 doi 10 1525 bio 2013 63 4 7 a b c Bockmann F A amp R M C Castro 2010 The blind catfish from the caves of Chapada Diamantina Bahia Brazil Siluriformes Heptapteridae description anatomy phylogenetic relationships natural history and biogeography Neotropical Ichthyology 8 4 673 706 doi 10 1590 s1679 62252010000400001 a b Schaefer Provenzano De Pinna amp Baskin 2005 New and Noteworthy Venezuelan Glanapterygine Catfishes Siluriformes Trichomycteridae with Discussion of Their Biogeography and Psammophily American Museum Novitates 3496 1 27 doi 10 1206 0003 0082 2005 496 0001 nanvgc 2 0 co 2 hdl 2246 5665 S2CID 19506818 Uiblein F Ott J A and Stachowitsch M 1996 Deep sea and extreme shallow water habitats Affinities and Adaptations Biosystematics and Ecology Series Band 11 ISBN 978 3 7001 2574 7 Lucanus Oliver 2013 First Notes on the Husbandry of the Blind Cichlid Lamprologus lethops from the Congo River Cichlid News vol 22 1 6 11 Flammang B E A Suvarnaraksha J Markiewicz amp D Soares 2016 Tetrapod like pelvic girdle in a walking cavefish Scientific Reports 6 23711 Bibcode 2016NatSR 623711F doi 10 1038 srep23711 PMC 4806330 PMID 27010864 a b Harries D T Arbenz N Dahanukar R Raghavan M Tringham D Rangad G Proudlove 2019 The world s largest known subterranean fish a discovery in Meghalaya NE India of a cave adapted fish related to the Golden Mahseer Tor putitora Hamilton 1822 Cave and Karst Science 46 3 121 126 a b Main D 12 February 2020 World s largest cave fish discovered in India National Geographic Retrieved 27 June 2020 a b c Muriel Cunha Janice de Pinna Mario 2005 New data on Cistern Catfish Phreatobius cisternarum from subterranean waters at the mouth of the Amazon River Siluriformes Incertae Sedis PDF Papeis Avulsos de Zoologia 35 327 339 Dutchen S 13 July 2015 Fat fish illuminate human obesity ScienceDaily Harvard Medical School Retrieved 26 April 2017 Armbruster J W M L Niemiller amp P B Hart 2016 Morphological Evolution of the Cave Spring and Swampfishes of the Amblyopsidae Percopsiformes Copeia 104 3 763 777 doi 10 1643 ci 15 339 S2CID 53608365 a b Moreira C R Bichuette M E Oyakawa O T de Pinna M C C and Trajano E 2010 Rediscovery and redescription of the unusual subterranean characiform Stygichthys typhlops with notes on its life history Journal of Fish Biology London Wiley InterScience 76 7 1815 1824 Vidthayanon C 2011 Cryptotora thamicola IUCN Red List of Threatened Species 2011 e T41407A10459372 doi 10 2305 IUCN UK 2011 1 RLTS T41407A10459372 en Retrieved 23 December 2017 a b c Poulson T L amp W B White 1969 The cave environment Science 165 3897 971 981 Bibcode 1969Sci 165 971P doi 10 1126 science 165 3897 971 PMID 17791021 McDowell I 10 November 2016 Alabama Cavefish Encyclopedia of Alabama Retrieved 16 May 2017 Nielsen Schwarzhans amp Hadiaty 2009 A blind new species of Diancistrus Teleostei Bythitidae from three caves on Muna Island southeast of Sulawesi Indonesia Cybium 33 3 241 245 Moller Schwarzhans Iliffe amp Nielsen 2006 Revision of the Bahamian cave fishes of the genus Lucifuga Ophidiiformes Bythitidae with description of a new species from islands on the Little Bahama Bank Zootaxa 33 1223 23 46 doi 10 11646 zootaxa 1223 1 3 Froese Rainer Pauly Daniel eds 2017 Milyeringa veritas in FishBase April 2017 version Ophisternon candidum Blind Cave Eel Department of the Environment Australia Retrieved 28 April 2017 a b Ma L and Y H Zhao 2012 Cavefish of China Pp 107 125 in White W B and D C Cuvier editors Encyclopedia of Caves Elsevier ISBN 9780123838322 a b Andy Coghlan 3 April 2017 First ever cavefish discovered in Europe evolved super fast New Scientist Retrieved 4 April 2017 a b c Proudlove G S 2010 Biodiversity and distribution of the subterranean fishes of the world Pp 41 63 in Trajano E Bichuette M E Kapoor B G eds The Biology of Subterranean Fishes Science ISBN 978 1578086702 Lina M Mesa S Carlos A Lasso Luz E Ochoa Carlos DoNascimiento 2018 Trichomycterus rosablanca Siluriformes Trichomycteridae a new species of hipogean catfish from the Colombian Andes Biota Colombiana 19 1 95 116 doi 10 21068 c2018 v19s1a09 Nguyen Dinh Tao Liang Cao Shuqing Deng E Zhang 2018 Speolabeo hokhanhi A New Cavefish from Central Vietnam Teleostei Cyprinidae Zootaxa 4476 1 109 117 doi 10 11646 zootaxa 4476 1 10 PMID 30313345 a b Chakrabarty Prosanta Prejean Jacques A Niemiller Matthew L 29 May 2014 The Hoosier cavefish a new and endangered species Amblyopsidae Amblyopsis from the caves of southern Indiana ZooKeys 412 41 57 doi 10 3897 zookeys 412 7245 PMC 4042695 PMID 24899861 Walsh S J Chakrabarty P 2016 A new genus and species of blind sleeper Teleostei Eleotridae from Oaxaca Mexico First obligate cave gobiiform in the Western Hemisphere Copeia 104 2 506 517 doi 10 1643 ci 15 275 S2CID 89252631 a b Proudlove G 2006 Subterranean fishes of the world ISBN 978 2 9527084 0 1 a b Dahanukar Neelesh Sundar Remya L Rangad Duwaki Proudlove Graham Raghavan Rajeev 2 June 2023 The world s largest cave fish from Meghalaya Northeast India is a new species Neolissochilus pnar Cyprinidae Torinae Vertebrate Zoology 73 141 152 doi 10 3897 vz 73 e101011 ISSN 2625 8498 Britz Ralf Kakkassery Francy Raghavan Rajeev 2014 Osteology of Kryptoglanis shajii a stygobitic catfish Teleostei Siluriformes from Peninsular India with a diagnosis of the new family Kryptoglanidae Ichthyological Exploration of Freshwaters 24 3 193 207 a b Raghavan Rajeev Dahanukar Neelesh Anoop V K Britz Ralf 2019 The subterranean Aenigmachanna gollum a new genus and species of snakehead Teleostei Channidae from Kerala South India Zootaxa 4603 2 377 388 doi 10 11646 zootaxa 4603 2 10 PMID 31717234 S2CID 164781147 a b Ravi Charan Basheer V S Kumar Rahul G 17 July 2019 Aenigmachanna mahabali a new species of troglophilic snakehead Pisces Channidae from Kerala India Zootaxa 4638 3 410 418 doi 10 11646 zootaxa 4638 3 6 ISSN 1175 5334 PMID 31712470 S2CID 203899040 Britz R 2016 Pillaiabrachia siniae a new species of earthworm eel from northern Myanmar Teleostei Synbranchiformes Chaudhuriidae Ichthyol Explor Freshwaters 27 1 41 47 Villa Verde Lima Carvalho amp Lima 2013 Rediscovery taxonomic and conservation status of the threatened catfish Listrura camposi Miranda Ribeiro Siluriformes Trichomycteridae Neotrop Ichthyol 11 1 55 64 doi 10 1590 S1679 62252013000100006 a b c d Romero Zhao amp Chen 2009 The Hypogean fishes of China Environ Biol Fish 86 211 278 doi 10 1007 s10641 009 9441 3 S2CID 41778476 University of Texas at Austin 17 June 2016 Rare blind catfish never before found in US discovered in national park cave in Texas ScienceDaily Retrieved 13 May 2017 Ohara W M I D Da Costa M L Fonseca 2016 Behavioiur feeding habits and ecology of the blind catfish Phreatobius sanguijuela Ostariophysi Siluriformes Journal of Fish Biology 89 2 1285 1301 doi 10 1111 jfb 13037 PMID 27329067 Espinasa Rivas Manzano amp Espinosa Perez 2001 A New Blind Cave Fish Population of Genus Astyanax Geography Morphology and Behavior Environmental Biology of Fishes 62 1 339 344 doi 10 1023 A 1011852603162 S2CID 30720408 Jeffery Strickler amp Yamamoto 2003 To See or Not to See Evolution of Eye Degeneration in Mexican Blind Cavefish Integr Comp Biol 43 4 531 541 doi 10 1093 icb 43 4 531 PMID 21680461 a b c Keene Yoshizawa amp McGaugh 2016 Biology and Evolution of the Mexican Cavefish pp 68 69 77 87 ISBN 978 0 12 802148 4 Zhang C G Zhao Y H 2016 Species Diversity and Distribution of Inland Fishes in China Science Press Beijing China ISBN 9787030472106 Nuryanto A D Bhagawati M N Abulias Indarmawan 2016 Ichtyofauna at Cijalu River Cilacap regency central Java Province Indonesia Biotropia 23 1 1 9 doi 10 11598 btb 2016 23 1 362 Kottelat M T Whitten 1996 Freshwater Biodiversity in Asia With Special Reference to Fish Vol 23 343 The World Bank p 32 via World Bank Technical Papers Proudlove G S 2019 Non stygobitic fishes in caves and other subterranean habitats Subterranean Fishes of the World Retrieved 17 January 2020 Farashi A Kaboli M Rezaei H R Naghavi M R Rahimian H Coad B W 2014 Reassessment of the taxonomic position of Iranocypris typhlops Bruun amp Kaiser 1944 Actinopterygii Cyprinidae ZooKeys 374 69 77 doi 10 3897 zookeys 374 6617 PMC 3909813 PMID 24493966 Hamidan N H M F Geiger J Freyhof 2014 Garra jordanica a new species from the Dead Sea basin with remarks on the relationship of G ghorensis G tibanica and G rufa Teleostei Cyprinidae Ichthyol Explor Freshwaters 25 3 223 236 Esmaeli H R G Sayyadzadeh B W Coad S Eagderi Review of the genus Garra Hamilton 1822 in Iran with description of a new species a morpho molecular approach Teleostei Cyprinidae Iran J Ichthyol 3 2 82 121 Zhang C amp Zhao Y H 2016 Species Diversity and Distribution of Inland Fishes in China Science Press p 296 ISBN 9787030472106 Kottelat M 2017 Speolabeo a new genus name for the cave fish Bangana musaei Teleostei Cyprinidae Zootaxa 4254 4 531 541 doi 10 11646 zootaxa 4254 4 6 PMID 28609956 Freyhof J E Baycelebi M Geiger 2018 Review of the genus Cobitis in the Middle East with the description of eight new species Teleostei Cobitidae Zootaxa 4535 1 1 75 doi 10 11646 zootaxa 4535 1 1 PMID 30647339 S2CID 58634705 a b c d Kottelat M 2012 Conspectus cobitidum an inventory of the loaches of the world Teleostei Cypriniformes Cobitoidei Raffles Bulletin of Zoology 26 1 199 doi 10 1007 s10641 009 9441 3 S2CID 41778476 Kottelat M 2010 Claea a new replacement name for Oreias Sauvage 1874 Teleostei Nemacheilidae Ichthyol Explor Freshwaters 21 4 384 Segherloo Ghaedrahmati amp Freyhof 2016 Eidinemacheilus a new generic name for Noemacheilus smithi Greenwood Teleostei Nemacheilidae Zootaxa 4147 4 466 476 doi 10 11646 zootaxa 4147 4 7 PMID 27515629 Shaji C P 2011 Indoreonectes evezardi IUCN Red List of Threatened Species 2011 e T10823A3219098 doi 10 2305 IUCN UK 2011 1 RLTS T10823A3219098 en Retrieved 23 December 2017 Tencatt amp Bichuette 2017 Aspidoras mephisto new species The first troglobitic Callichthyidae Teleostei Siluriformes from South America PLOS ONE 12 3 e0171309 Bibcode 2017PLoSO 1271309T doi 10 1371 journal pone 0171309 PMC 5331963 PMID 28248959 Froese Rainer Pauly Daniel eds 2017 Rhamdia laticauda in FishBase May 2017 version Froese Rainer Pauly Daniel eds 2017 Rhamdia quelen in FishBase May 2017 version Binoy Roshan amp Rakesh 2012 Occurrence of Kryptoglanis shajii an enigmatic subterranean spring catfish Siluriformes Incertae sedis in the channels of paddy fields Current Science 102 2 161 Froese Rainer Pauly Daniel eds 2017 Forbesichthys agassizii in FishBase May 2017 version Niemiller Near amp Fitzpatrick 2011 Delimiting species using multilocus data diagnosing cryptic diversity in the southern cavefish Typhlichthys subterraneus Teleostei Amblyopsidae Evolution 66 3 846 866 doi 10 1111 j 1558 5646 2011 01480 x PMID 22380444 S2CID 7790397 Espinasa L amp W R Jeffery 2003 A troglomorphic sculpin Pisces Cottidae population geography morphology and conservation status Journal of Cave and Karst Studies 65 2 93 100 Williams J D amp W M Howell 1979 An albino sculpin from a cave in the New River drainage of West Virginia Pisces Cottidae Brimleyana 1 141 146 a b Adams G L B M Burr J L Day amp D E Starkey 2013 Cottus specus a new troglomorphic species of sculpin Cottidae from southeastern Missouri Zootaxa 3609 5 484 494 doi 10 11646 zootaxa 3609 5 4 PMID 24699612 Pouyaud Kadarusman Hadiaty Slembrouck Lemauk Kusumah amp Keith 2013 Oxyeleotris colasi Teleostei Eleotridae a new blind cave fish from Lengguru in West Papua Indonesia Cybium 36 4 521 529 a b c Chakrabarty P 2010 Status and phylogeny of Milyeringidae Teleostei Gobiiformes with the description of a new blind cave fish from Australia Milyeringa brooksi n sp Zootaxa 2557 19 28 doi 10 11646 zootaxa 2557 1 2 a b c Sparks J S amp P Chakrabarty 2012 Revision of the endemic Malagasy Cavefish genus Typhleotris Teleostei Gobiiformes Milyeringidae with discussion of its phylogenetic placement and description of a new species American Museum Novitates 3764 1 28 doi 10 1206 3764 2 hdl 2246 6399 S2CID 85731146 NatureServe 2013 Speoplatyrhinus poulsoni IUCN Red List of Threatened Species 2013 e T20467A19033986 doi 10 2305 IUCN UK 2013 1 RLTS T20467A19033986 en Retrieved 23 December 2017 Bruton M N 1995 Threatened fishes of the world Clarias cavernicola Trewavas 1936 Clariidae Environmental Biology of Fishes 43 2 162 doi 10 1007 BF00002486 S2CID 44350023 Freyhof J 2014 Caecocypris basimi IUCN Red List of Threatened Species 2014 e T3450A19006223 doi 10 2305 IUCN UK 2014 1 RLTS T3450A19006223 en Retrieved 23 December 2017 Montanari M 30 June 2016 This Rare Eyeless Cavefish Was Discovered Deep Underground In Mexico Forbes Retrieved 30 April 2017 Alabama Cavefish U S Fish and Wildlife Service Archived from the original on 7 October 2014 Retrieved 12 October 2011 Klaus S amp M Plath 2011 Predation on cave fish by freshwater crab Avotrichodactylus bidens Bott 1969 Brachyura Trichodactylidae in Mexican sulfur cave Crustaceana 84 4 411 418 doi 10 1163 001121611X560853 Tobler M 2009 Does a predatory insect contribute to the divergence between cave and surface adapted fish populations Biol Lett 5 4 506 509 doi 10 1098 rsbl 2009 0272 PMC 2781934 PMID 19443506 Horstkotte Riesch Plath amp Jager 2010 Predation by three species of spiders on a cave fish in a Mexican sulphur cave Bull Br Arachnol Soc 15 2 55 58 doi 10 13156 arac 2010 15 2 55 S2CID 41990323 SeriouslyFish Astyanax mexicanus Retrieved 28 February 2016 Romero A 2009 Cave Biology Life in Darkness Cambridge University Press pp 147 148 ISBN 978 0 521 82846 8 Retrieved from https en wikipedia org w index php title Cavefish amp oldid 1142947980, wikipedia, wiki, book, books, library,

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