Cyprinodon

Last updated

Contents

Cyprinodon
Owens pupfish (Cyprinodon radiosus).jpg
Cyprinodon radiosus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Cyprinodontiformes
Family: Cyprinodontidae
Subfamily: Cyprinodontinae
Genus: Cyprinodon
Lacépède, 1803
Type species
Cyprinodon variegatus
Lacépède, 1803
Synonyms
  • EncratesGistel, 1848
  • LebiaOken, 1817
  • PrinodonRafinesque, 1815
  • TrifarciusPoey, 1860

Cyprinodon is a genus of pupfishes found in waters that range from fresh to hypersaline. The genus is primarily found in Mexico, the Caribbean Islands and southern United States (Arizona, California, Florida, Nevada, New Mexico, Oklahoma and Texas), but C. variegatus occurs as far north as Massachusetts and along the entire Gulf of Mexico coastline, and C. dearborni and C. variegatus are found in northern South America. [1] [2] Many species have tiny ranges and are highly threatened, in some cases already extinct. [2] [3] [4] Cyprinodon are small; the largest reaches 10 cm (3.9 in) in length and most other species only reach about half that size. [1]

Distribution and habitat

Cyprinodon variegatus is the most widespread member of the genus. Cyprinodon variegatus (S0103) (12598764423).jpg
Cyprinodon variegatus is the most widespread member of the genus.
Cyprinodon diabolis is very rare; its entire native range is the Devils Hole in Nevada Cyprinodon diabolis Wales, 1930.jpg
Cyprinodon diabolis is very rare; its entire native range is the Devils Hole in Nevada

A few Cyprinodon species have quite large ranges, notably C. variegatus , but the vast majority have small ranges, typically restricted to one or two Mexican states or US states, Hispaniola, or a Bahaman island. C. longidorsalis and C. diabolis have both been said to have the smallest native range of any vertebrate species, with the former being restricted to a spring pool that covers about 10 m2 (110 sq ft) and the latter to an 18 m2 (190 sq ft) shelf in a spring pool, but C. longidorsalis is now only found in captivity as its habitat has disappeared. [3] [5] While most Cyprinodon species have separate distributions, seven (C. beltrani, C. esconditus, C. labiosus, C. maya, C. simus, C. suavium and C. verecundus) are endemic in Lake Chichancanab in Quintana Roo, Mexico, [6] [7] and three (the endemic C. brontotheroides and C. desquamator, and the widespread C. variegatus) live in the hypersaline lakes on San Salvador Island in the Bahamas. [2] In a few other cases separate species do come into contact towards the edges of their distributions where they often hybridize, notably C. eximius X C. pachycephalus and C. atrorus X C. bifasciatus, but also C. variegatus in places where it has been introduced by humans into the ranges of other Cyprinodon species. [2]

Although the individual Cyprinodon species often have a highly specific habitat, overall the genus occurs in a remarkable range of places, such as springs (including those isolated in deserts), pools, lakes, coastal lagoons, creeks, streams and rivers. Their salinity and temperature range is very broad, like those living in hot springs (taken to the extreme in C. julimes in water up to 46 °C or 115 °F, and C. pachycephalus up to 49 °C or 120 °F), [8] [9] and those of hypersaline habitats where the salinity far exceeds that of sea water. [10] [11] Certain species may even experience very large variations in the temperature and salinity over a relatively short period. For example, some populations of C. variegatus live in water where the temperature has been known to change from 15 to −1.8 °C (59 to 29 °F) in less than 24 hours (in the coldest temperatures they bury into the substrate). [12] Some populations of C. nevadensis tolerate water temperatures between 2 and 44 °C (36–111 °F), and C. salinus live in waters where the temperature may change by as much as 19 °C (34 °F) in a day and 40 °C (72 °F) in a season. [12] [13] Furthermore, the salinity of C. salinus' habitat may vary from less than one-third of that of sea water to almost five times as much as sea water in a season. [10]

Conservation status

Most species in the genus are seriously threatened. C. arcuatus, C. ceciliae, C. inmemoriam, C. nevadensis calidae and an undescribed species popularly known as the "Perrito de Sandia" are already extinct. [13] [14] [15] [16] [17] C. arcuatus was restricted to springs in the US state of Arizona and probably also in the Mexican state of Sonora, [14] while all the others were restricted to spring systems in Mexico. [18] [19]

Three species from southwestern Nuevo León, C. alvarezi, C. longidorsalis and C. veronicae, have become extinct in the wild, only surviving in captivity. [18] [20] [21] A few other Mexican species still considered endangered or vulnerable by the IUCN, including at least C. maya, C. simus and C. verecundus of Lake Chichancanab, also appear to only survive in captivity. [22] [23] Several others have very small remaining populations in the wild. [19] Among the species that survive in the wild, the rarest is perhaps C. diabolis from the tiny Devils Hole in Nevada; in recent decades its population has fluctuated between a few tens and a few hundred individuals. [24] Primary threats to pupfish are habitat loss due to water extraction, drought and pollution, and introduced species. [3] [19]

Behavior

Feeding

Cyprinodon desquamator, the only known scale-eating species of pupfish Cyprinodon Desquamator.jpg
Cyprinodon desquamator , the only known scale-eating species of pupfish

Most Cyprinodon species feed on algae, cyanobacteria and detritus, but may also supplement their diet with small crustaceans and aquatic insect larvae. [13] [22] Some species mainly feed on small animals like aquatic insects. [11] C. variegatus, a species that otherwise has a diet typical of pupfish, will clean other fish by feeding on parasites on their body. [25]

In the two places where several species live together they have diverged into different niches, including the fish-eating C. maya (Lake Chichancanab), zooplankton-eating C. simus (Lake Chichancanab), amphipod- and bivalve-eating C. labiosus and C. verecundus (Lake Chichancanab), scale-eating C. desquamator (San Salvador Island lakes), and ostracod- and gastropod-eating C. brontotheroides (San Salvador Island lakes). [2] [22]

Breeding

Macularius stimulation.jpg
Salt Creek pupfish.jpg
On left: Cyprinodon macularius pair during courtship (male above).
On right: Cyprinodon salinus pair during spawning (male with yellowish back).

Cyprinodon pupfish are short-lived, typically reaching an age of no more than a year in the wild, [13] although some may reach as much as three years. [25] Especially those from habitats that experience major environmental fluctuations (for example, large variations in temperature) rapidly reach maturity and are already able to breed when 1–1.5 month old. [13] Despite some species' ability to survive in a wide temperature range, their requirements for breeding often are much more specific. For example, although some populations of C. nevadensis can live in water that ranges between 2 and 44 °C (36–111 °F), they only breed from 24 to 30 °C (75–86 °F). [13] However, there are exceptions like C. rubrofluviatilis that will breed at a relative wide range from 13 to 34 °C (55–93 °F). [11] Consequently, pupfish living in stable habitats breed year-round, but those in more seasonal habitats generally only at certain times of the year where the conditions are optimal. [13] When breeding, males assume a relatively bright nuptial coloration. [13]

There are two primary breeding strategies: In species of small isolated habitats like springs, each large male (or medium-sized male, if large males are absent) defends a territory and displays to visiting females that will lay their eggs inside the territory. [13] [26] In at least some species, small males will attempt to fertilize eggs by sneaking into a territory of a larger male. [26] Once deposited, neither sex cares for the eggs, although they do get a level of protection by being inside the territory of a male. A male will attempt to attract several females to lay their eggs in his territory and a female may lay eggs in the territories of several males. [13] Another breeding strategy is used by species that inhabit rivers. Here the males do not maintain a territory and groups of pupfish gather to breed. A male typically will lead a female to the edge of the group to spawn, although on occasion it may occur in the middle of the group. [13] The eggs of Cyprinodon pupfish are adhesive and stick to the substrate, [13] or they are covered in sand. [25]

Species

Cyprinodon brontotheroides (shown) and C. desquamator are both restricted to hypersaline lakes in the Bahamas Cyprinodon brontotheroides.jpg
Cyprinodon brontotheroides (shown) and C. desquamator are both restricted to hypersaline lakes in the Bahamas
Cyprinodon eremus in shallow water, which is preferred by most species in this genus Sonoyta pupfish.jpg
Cyprinodon eremus in shallow water, which is preferred by most species in this genus
Cyprinodon julimes was only scientifically described in 2009 Julimes pupfish (Cyprinodon julimes) male.jpg
Cyprinodon julimes was only scientifically described in 2009
Cyprinodon rubrofluviatilis, a relatively common species from Texas Cyprinodon rubrofluviatilis.jpg
Cyprinodon rubrofluviatilis , a relatively common species from Texas
Cyprinodon desquamator (shown), the scale-eating pupfish, in its natural habitat on San Salvador Island, Bahamas Cyprinodon desquamator.jpg
Cyprinodon desquamator (shown), the scale-eating pupfish, in its natural habitat on San Salvador Island, Bahamas
Cyprinodon macrolepis male in an aquarium. Cyprinodon macrolepis.tif
Cyprinodon macrolepis male in an aquarium.

There are currently 49 recognized species in this genus: [1] [2]

Related Research Articles

<span class="mw-page-title-main">Death Valley pupfish</span> Small endangered fish native to Death Valley, California

The Death Valley pupfish, also known as Salt Creek pupfish, is a small species of fish in the family Cyprinodontidae found only in Death Valley National Park, California, United States. There are two recognized subspecies: C. s. salinus and C. s. milleri. The Death Valley pupfish is endemic to two small, isolated locations and currently classified as endangered.

<span class="mw-page-title-main">Tecopa pupfish</span> Extinct subspecies of fish

The Tecopa pupfish is an extinct subspecies of the Amargosa pupfish. The small, heat-tolerant pupfish was endemic to the outflows of a pair of hot springs in the Mojave Desert of Inyo County, California. Habitat modifications, the introduction of non-native species and hybridization with the related Amargosa River pupfish led to its extinction around 1979.

The Shoshone pupfish is a subspecies of Cyprinodon nevadensis from California in the United States. It is characterized by large scales and a "slab-sided," narrow, slender body, with the arch of the ventral contour much less pronounced than the dorsal. It also has fewer pelvic fin rays and scales than the other subspecies of C. nevadensis.

Robert Rush Miller was an important figure in American ichthyology and conservation from 1940 to the 1990s.

Potosi pupfish is a species of fish in the family Cyprinodontidae. It is endemic to Mexico, but is now extinct in the wild, only surviving in captivity.

Blackfin pupfish is a species of fish in the family Cyprinodontidae. This pupfish is endemic to Lake Chichancanab in Quintana Roo, Mexico. In almost all places, different Cyprinodon species do not overlap in their range, but there are two notable exceptions and one of these is Lake Chichancanab, which is inhabited by C. beltrani, C. esconditus, C. labiosus, C. maya, C. simus, C. suavium and C. verecundus. Among the endemic Cyprinodon species in Lake Chichancanab, only C. beltrani and C. labiosus still occur in some numbers in their habitat, while the remaining are virtually—if not fully—extinct in the wild. This species feeds on detritus. Compared to the other species of pupfish mentioned previously, the C. beltrani prefer a different feeding substrate. In regards to mating, the female blackfin pupfish seems to mate with other species of pupfish and not just the C. beltrani. The male blackfin pupfish have similar markings and courting behaviors to other pupfish in their habitat. The person honored in the specific name of this pupfish is the Mexican biologist Enrique Beltrán Castillo (1903–1994) to mark his quarter century of being a biologist.

Cyprinodon ceciliae is an extinct species of pupfish. It was endemic to the Ojo de Agua la Presa in southwestern Nuevo Leon state in Mexico, but disappeared in 1990 due to habitat loss. The same freshwater spring system was the home of three other pupfish: Cyprinodon inmemoriam (extinct), Cyprinodon longidorsalis and Cyprinodon veronicae. Although these were from the same spring system, each was restricted to its own individual spring and associated waters. The exact spring and associated waters inhabited by the Villa Lopez pupfish were also the home of a now-extinct, undescribed species of Cambarellus crayfish. The specific name honours the daughter of María de Lourdes Lozano-Vilano and niece of Salvador Contreras-Balderas, Cecilia Contreras Lozano, who assisted on the trip on which the type was collected.

The Cachorrito de la Trinidad, also known in English as the Charco Azul pupfish, was a small species of fish in the family Cyprinodontidae. It was endemic to Ojo de Agua la Presa in southwestern Nuevo Leon state in Mexico, but became extinct in 1985 due to habitat loss. The same freshwater spring system was the home of three other pupfish: Cyprinodon ceciliae (extinct), Cyprinodon longidorsalis and Cyprinodon veronicae. Although these were from the same spring system, each was restricted to its own individual spring pool. The spring pond inhabited by the Charco Azul pupfish was also the home of a now-extinct, undescribed species of Cambarellus crayfish. The specific name inmemoriam means "in memory" and signifies the extinction of the species due to its native spring being dried out by groundwater extraction soon after the discovery of this fish in 1983.

The thicklip pupfish, known in Spanish as Cachorrito cangrejero, is a species of fish in the family Cyprinodontidae. The thicklip pupfish is endemic to Laguna Chichancanab in Quintana Roo, Mexico. In almost all places, different Cyprinodon species do not overlap in their range, but there are two notable exceptions and one of these is Lake Chichancanab, which is inhabited by C. labiosus, C. beltrani, C. esconditus, C. maya, C. simus, C. suavium and C. verecundus. Among the endemic Cyprinodon species in Lake Chichancanab, only C. beltrani and C. labiosus still occur in some numbers in their habitat, while the remaining are virtually—if not fully—extinct in the wild. They usually live in a tropical climate with temperatures ranging from 22 to 26 °C. The maximum total length of this fish is 7 centimetres (2.8 in).

Cyprinodon longidorsalis, the cachorrito de charco palmal or La Palma pupfish, is a species of fish in the family Cyprinodontidae. It was endemic to the Ojo de Agua la Presa in southwestern Nuevo Leon state in Mexico, but became extinct in the wild in 1994 due to habitat loss. The same freshwater spring system was the home of three other pupfish: Cyprinodon ceciliae (extinct), Cyprinodon inmemoriam (extinct) and Cyprinodon veronicae. Although these were from the same spring system, each was restricted to its own individual spring pool. The Charco La Palma pool and its spring had a combined area of about 10 m2 (110 sq ft) and was no more than 1.4 m at the deepest point, making the range of the La Palma pupfish perhaps the smallest known for any vertebrate species. This tiny spring pond also was the home of a now-extinct, undescribed species of Cambarellus crayfish.

The Maya pupfish, known in Spanish as cachorrito gigante, is a highly threatened species of fish in the family Cyprinodontidae. It is endemic to Lake Chichancanab in Quintana Roo, Mexico. In almost all places, different Cyprinodon species do not overlap in their range, but there are two notable exceptions and one of these is Lake Chichancanab, which is inhabited by C. maya, C. beltrani, C. esconditus, C. labiosus, C. simus, C. suavium and C. verecundus. Living together, the Cyprinodon species in Lake Chichancanab have diverged into different niches. Pupfish typically feed on algae and detritus. In Lake Chichancanab, however, C. maya has become not only the largest species in the genus Cyprinodon, up to 10 cm (3.9 in) long, but also the only that catches and eats whole fish. In smaller quantities it eats ostracods and freshwater snails.

<span class="mw-page-title-main">Pecos pupfish</span> Species of fish

The Pecos pupfish is a species of pupfish in the family Cyprinodontidae. It is endemic to the Pecos River watershed in eastern New Mexico and western Texas in the United States.

<span class="mw-page-title-main">Owens pupfish</span> Species of fish

The Owens pupfish is a rare species of fish in the family Cyprinodontidae, the pupfish. It is endemic to California in the United States, where it is limited to the Owens Valley. It is a federally listed endangered species of the United States. This pupfish is up to 5 centimetres long, the largest males sometimes longer. The male is blue-gray, turning bright blue during spawning. The female is greenish brown with a silvery or whitish belly.

The boxer pupfish is a small species of pupfish in the family Cyprinodontidae. It is endemic to Lake Chichancanab in Quintana Roo, Mexico.

The largefin pupfish, also known as cachorrito de dorsal larga, is a small species of pupfish in the family Cyprinodontidae. It is endemic to Lake Chichancanab in Quintana Roo, Mexico. In almost all places, different Cyprinodon species do not overlap in their range, but there are two notable exceptions and one of these is Lake Chichancanab, which is inhabited by C. verecundus, C. beltrani, C. esconditus, C. labiosus, C. maya, C. simus and C. suavium. Living together, the Cyprinodon species in Lake Chichancanab have diverged into different niches. Pupfish typically feed on algae and detritus. In Lake Chichancanab, however, C. verecundus has become an amphipod- and bivalve-eater.

The Charco Palma pupfish is a species of small fish in the family Cyprinodontidae. It is endemic to the Ojo de Agua la Presa in southwestern Nuevo Leon state in Mexico. Although listed as critically endangered by the IUCN in 1996, the species is now extinct in the wild. The same freshwater spring system was the home of three other pupfish: Cyprinodon ceciliae (extinct), Cyprinodon inmemoriam (extinct) and Cyprinodon longidorsalis. Although these were from the same spring system, each was restricted to its own individual spring and associated waters. The exact spring and associated waters inhabited by the Charco Palma pupfish were also the home of three now-extinct invertebrates: An undescribed species of Cambarellus crayfish, the valvatid freshwater snail Valvata beltrani and an undescribed species of valvatid freshwater snail. The specific name honours the niece of María de Lourdes Lozano-Vilano and daughter of Salvador Contreras-Balderas, Verónica Contreras Arqueita, who assisted on the trip on which the type was collected.

<span class="mw-page-title-main">Catarina pupfish</span> Species of fish

The Catarina pupfish was a diminutive species of fish in the family Cyprinodontidae, first described in 1972. It was endemic to a spring in Nuevo León, Mexico. In an attempt of saving the rapidly declining species, some were brought into captivity in the late 1980s and early 1990s, but it proved very difficult to maintain. In 1994 it became extinct in the wild. Gradually the captive populations also perished. The last male died in 2014 and the species became extinct.

<span class="mw-page-title-main">Desert pupfish</span> Species of fish

The desert pupfish is a rare species of bony fish in the family Cyprinodontidae. It is a small fish, typically less than 7.62 cm (3 in) in length. Males are generally larger than females, and have bright-blue coloration, while females and juveniles are silvery or tan. A notable attribute of the desert pupfish is their ability to survive in environments of extreme salinity, pH, and temperature, and low oxygen content. The desert pupfish mates in a characteristic fashion, wherein compatible males and females will come in contact and collectively jerk in an s-shape. Each jerk typically produces a single egg that is fertilized by the male and deposited in his territory. Breeding behavior includes aggressive arena-breeding and more docile consort-pair breeding.

<span class="mw-page-title-main">Sonoyta pupfish</span> Species of fish

The Sonoyta pupfish or Quitobaquito pupfish is an endangered species of pupfish from Sonora in Mexico and Arizona in the United States.

<i>Cyprinodon julimes</i> Species of fish

The Julimes pupfish is a species of killifish in the family Cyprinodontidae. This pupfish is endemic to El Pandeño, a hot spring in Julimes, Chihuahua, Mexico, and it is adapted to life in water that reaches temperatures as high as 46 °C (114 °F). Consequently, it has been referred to as the "hottest fish in the world", although the closely related Cyprinodon pachycephalus can occur in somewhat warmer waters. Cyprinodon julimes was scientifically described in 2009 as similar to Cyprinodon eximius, but it has a bigger head, nearly one-third of its standard length.

References

  1. 1 2 3 Froese, Rainer and Pauly, Daniel, eds. (2018). Species of Cyprinodon in FishBase . October 2018 version.
  2. 1 2 3 4 5 6 7 8 Martin, C. H., and P. C. Wainwright (2013). Multiple Fitness Peaks on the Adaptive Landscape Drive Adaptive Radiation in the Wild. Science 339(6116): 208-211.
  3. 1 2 3 Contreras-Balderas, S.; L. Lozano-Vilano (1996). "Extinction of most Sandia and Potosí valleys (Nuevo León, Mexico) endemic pupfishes, crayfishes and snails". Ichthyol. Explor. Freshwaters. 7 (1): 33–40.
  4. Martin, Christopher H.; Crawford, Jacob E.; Turner, Bruce J.; Simons, Lee H. (2016-01-27). "Diabolical survival in Death Valley: recent pupfish colonization, gene flow and genetic assimilation in the smallest species range on earth". Proceedings of the Royal Society B: Biological Sciences. 283 (1823): 20152334. doi: 10.1098/rspb.2015.2334 . ISSN   0962-8452. PMC   4795021 . PMID   26817777.
  5. Helfman, G.; B.B. Collette; D.E. Facey; B.W. Bowen (2009). The Diversity of Fishes: Biology, Evolution, and Ecology . Wiley-Blackwell. p.  413. ISBN   978-1-4051-2494-2.
  6. Contreras Balderas, S., and W. Bussing |(2013). Yucatan . Freshwater Ecoregions of the World. Retrieved 1 March 2013.
  7. Strecker, U. (2006). "The impact of invasive fish on an endemic Cyprinodon species flock (Teleostei) from Laguna Chichancanab, Yucatan, Mexico". Ecology of Freshwater Fish. 15 (4): 408–418. Bibcode:2006EcoFF..15..408S. doi: 10.1111/j.1600-0633.2006.00159.x .
  8. Clarke, A. (2017). Principles of Thermal Ecology: Temperature, Energy and Life. Oxford University Press. p. 323. ISBN   978-0-19-955166-8.
  9. Maiz-Tome, L. (2019). "Cyprinodon pachycephalus". IUCN Red List of Threatened Species . 2019: e.T6162A12524752. doi: 10.2305/IUCN.UK.1996.RLTS.T6162A12524752.en . Retrieved 17 October 2022.
  10. 1 2 Otto, R.G.; S.D. Gerking (1973). "Heat Tolerance of a Death Valley Pupfish (Genus Cyprinodon)". Physiological Zoology. 46 (1): 43–49. doi:10.1086/physzool.46.1.30152515. S2CID   87681268.
  11. 1 2 3 Hassan-Williams, C.; T.H. Bonner (2007). "Cyprinodon rubrofluviatilis". Texas Freshwater Fishes, Texas State University - San Marcos, Department of Biology. Retrieved 26 August 2019.
  12. 1 2 Bennett, W.A.; T.L. Beitinger (1997). "Temperature Tolerance of the Sheepshead Minnow, Cyprinodon variegatus". Copeia. 1997 (1): 77–87. doi:10.2307/1447842. JSTOR   1447842.
  13. 1 2 3 4 5 6 7 8 9 10 11 12 Moyle, P.B.; R.M. Yoshiyame; J.E. Williams; E.D. Wirkamanayake (1995). "Fish Species of Special Concern in California" (PDF). California Department of Fish and Game. Archived from the original (PDF) on 23 March 2012.
  14. 1 2 NatureServe (2013). "Cyprinodon arcuatus". IUCN Red List of Threatened Species . 2013: e.T202375A15362268. doi: 10.2305/IUCN.UK.2013-1.RLTS.T202375A15362268.en . Retrieved 12 November 2021.
  15. Contreras-Balderas, S.; Almada-Villela, P.; Palmer-Newton, A. (2019). "Cyprinodon ceciliae". IUCN Red List of Threatened Species . 2019: e.T6177A3107534. doi: 10.2305/IUCN.UK.2019-2.RLTS.T6177A3107534.en . Retrieved 12 November 2021.
  16. Schmitter-Soto, J. (2019). "Cyprinodon inmemoriam". IUCN Red List of Threatened Species . 2019: e.T6176A3107502. doi: 10.2305/IUCN.UK.2019-2.RLTS.T6176A3107502.en . Retrieved 12 November 2021.
  17. Contreras-Balderas, S.; Almada-Villela, P. (1996). "Cyprinodon spp.". IUCN Red List of Threatened Species . 1996. Retrieved 1 March 2013.
  18. 1 2 Contreras-MacBeath, T.; M.B. Rodríguez; V. Sorani; C. Goldspink; G.M. Reid (2014). "Richness and endemism of the freshwater fishes of Mexico". Journal of Threatened Taxa. 6 (2): 5421–5433. doi: 10.11609/JoTT.o3633.5421-33 .
  19. 1 2 3 Ceballos, G.; E.D. Pardo; L.M. Estévez; H.E. Pérez, eds. (2016). Los peces dulceacuícolas de México en peligro de extinción. Fondo de Cultura Económic. ISBN   978-607-16-4087-1.
  20. Valdes Gonzales, A. (2019). "Cyprinodon alvarezi". IUCN Red List of Threatened Species . 2019: e.T6144A3104750. doi: 10.2305/IUCN.UK.2019-2.RLTS.T6144A3104750.en . Retrieved 12 November 2021.
  21. Valdes Gonzales, A. (2019). "Cyprinodon longidorsalis". IUCN Red List of Threatened Species . 2019: e.T6174A3107266. doi: 10.2305/IUCN.UK.2019-2.RLTS.T6174A3107266.en . Retrieved 12 November 2021.
  22. 1 2 3 Martin, C.; P.C. Wainwright (2011). "Trophic novelty is linked to exceptional rates of morphological diversification in two adaptive radiations of Cyprinodon pupfish". Evolution. 65 (8): 2197–2212. doi: 10.1111/j.1558-5646.2011.01294.x . PMID   21790569.
  23. Martin, C.H.; Crawford, J.E.; Turner, B.J.; Simons, L.H. (2016). "Diabolical survival in Death Valley: recent pupfish colonization, gene flow and genetic assimilation in the smallest species range on earth". Proc Biol Sci. 283 (1823): 20152334. doi:10.1098/rspb.2015.2334. PMC   4795021 . PMID   26817777.
  24. "Devils Hole Pupfish". U.S. Fish and Wildlife Service. December 2, 2013. Retrieved 7 October 2018.
  25. 1 2 3 Hassan-Williams, C.; T.H. Bonner (2007). "Cyprinodon variegatus". Texas Freshwater Fishes, Texas State University - San Marcos, Department of Biology. Retrieved 26 August 2019.
  26. 1 2 Leiser, J.K.; M. Itzkowitz (2002). "The relative costs and benefits of territorial defense and the two conditional male mating tactics in the Comanche Springs pupfish (Cyprinodon elegans)". Acta Ethol. 5 (1): 65–72. doi:10.1007/s10211-002-0066-1. S2CID   42642403.
  27. Lyons, T.J. (2021). "Cyprinodon higuey". IUCN Red List of Threatened Species . 2021: e.T82958669A82962221. doi: 10.2305/IUCN.UK.2021-1.RLTS.T82958669A82962221.en . Retrieved 12 November 2021.
  28. Valdes Gonzales, A. (2019). "Cyprinodon latifasciatus". IUCN Red List of Threatened Species . 2019: e.T6155A3105057. doi: 10.2305/IUCN.UK.2019-2.RLTS.T6155A3105057.en . Retrieved 12 November 2021.