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Cyprinodon

{{Short description|Genus of fishes}}

{{Automatic taxobox

| fossil_range = {{fossil range|Late Miocene|present}}

| image = Owens pupfish (Cyprinodon radiosus).jpg

| image_caption = Cyprinodon radiosus

| taxon = Cyprinodon

| authority = Lacépède, 1803

| type_species = Cyprinodon variegatus

| type_species_authority = Lacépède, 1803

| synonyms = * Encrates Gistel, 1848

  • Lebia Oken, 1817
  • Prinodon Rafinesque, 1815
  • Trifarcius Poey, 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.{{FishBase genus | genus = Cyprinodon| month = October | year = 2018}}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. Many species have tiny ranges and are highly threatened, in some cases already extinct.{{cite journal| author1=Contreras-Balderas, S. | author2=L. Lozano-Vilano | year=1996| title=Extinction of most Sandia and Potosí valleys (Nuevo León, Mexico) endemic pupfishes, crayfishes and snails | journal=Ichthyol. Explor. Freshwaters | volume=7 | issue=1 | pages=33–40 }}{{Cite journal|last1=Martin|first1=Christopher H.|last2=Crawford|first2=Jacob E.|last3=Turner|first3=Bruce J.|last4=Simons|first4=Lee H.|date=2016-01-27|title=Diabolical survival in Death Valley: recent pupfish colonization, gene flow and genetic assimilation in the smallest species range on earth|journal=Proceedings of the Royal Society B: Biological Sciences|volume=283|issue=1823|pages=20152334|doi=10.1098/rspb.2015.2334|pmid=26817777|pmc=4795021|issn=0962-8452|doi-access=free}} Cyprinodon are small; the largest reaches {{convert|10|cm|abbr=on}} in length and most other species only reach about half that size.

Evolution

Based on phylogenetic evidence, Cyprinodon diverged from its closest relative, the recently extinct Megupsilon, during the Late Miocene, and saw a rapid evolutionary radiation afterwards. The only known fossil species from the genus is C. breviradius from the late Miocene or early Pliocene-aged sediments of Death Valley National Park (originally considered part of the Late Eocene-aged Titus Canyon Formation).{{Cite journal |last1=Echelle |first1=Anthony A. |last2=Carson |first2=Evan W. |last3=Echelle |first3=Alice F. |last4=Van Den Bussche |first4=R. A. |last5=Dowling |first5=Thomas E. |last6=Meyer |first6=Axel |date=2005 |editor-last=Wood |editor-first=R. M. |title=Historical Biogeography of the New-World Pupfish Genus Cyprinodon (Teleostei: Cyprinodontidae) |url=https://bioone.org/doi/abs/10.1643/CG-03-093R3 |journal=Copeia |language=en |volume=2005 |issue=2 |pages=320–339 |doi=10.1643/CG-03-093R3 |issn=0045-8511}}{{Cite web |title=PBDB Taxon |url=https://paleobiodb.org/classic/basicTaxonInfo?taxon_no=395089 |access-date=2025-03-20 |website=paleobiodb.org}}

Distribution and habitat

File:Cyprinodon variegatus (S0103) (12598764423).jpg is the most widespread member of the genus.]]

File:Cyprinodon diabolis Wales, 1930.jpg 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 U.S. 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 {{cvt|10|m2}} and the latter to an {{cvt|18|m2}} shelf in a spring pool, but C. longidorsalis is now only found in captivity as its habitat has disappeared.{{cite book | author1=Helfman, G. | author2=B.B. Collette | author3=D.E. Facey | author4=B.W. Bowen | year=2009 | title=The Diversity of Fishes: Biology, Evolution, and Ecology | url=https://archive.org/details/diversityfishesb00helf_991 | url-access=limited | page=[https://archive.org/details/diversityfishesb00helf_991/page/n430 413] | publisher=Wiley-Blackwell | isbn=978-1-4051-2494-2 }} 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,Contreras Balderas, S., and W. Bussing |(2013). [http://www.feow.org/ecoregions/details/175 Yucatan]. Freshwater Ecoregions of the World. Retrieved 1 March 2013.{{cite journal| author=Strecker, U. | year=2006 | title=The impact of invasive fish on an endemic Cyprinodon species flock (Teleostei) from Laguna Chichancanab, Yucatan, Mexico | journal=Ecology of Freshwater Fish | volume=15 | issue=4 | pages=408–418 | doi=10.1111/j.1600-0633.2006.00159.x | doi-access=free | bibcode=2006EcoFF..15..408S }} 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. 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.

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 {{cvt|46|C|disp=or}}, and C. pachycephalus up to {{cvt|49|C|disp=or}}),{{cite book | author=Clarke, A. | year=2017 | title=Principles of Thermal Ecology: Temperature, Energy and Life | page=323 | publisher=Oxford University Press | isbn=978-0-19-955166-8 }}{{Cite iucn | author = Maiz-Tome, L. | name-list-style = amp | title = Cyprinodon pachycephalus | volume = 2019 | page = e.T6162A12524752 | date = 2019 | doi = 10.2305/IUCN.UK.1996.RLTS.T6162A12524752.en|access-date=17 October 2022 }} and those of hypersaline habitats where the salinity far exceeds that of sea water.{{cite journal| author1=Otto, R.G. | author2=S.D. Gerking | year=1973 | title=Heat Tolerance of a Death Valley Pupfish (Genus Cyprinodon) | journal=Physiological Zoology | volume=46 | issue=1 | pages=43–49 | doi=10.1086/physzool.46.1.30152515| s2cid=87681268 }}{{cite web| author1=Hassan-Williams, C. | author2=T.H. Bonner | year=2007 | title=Cyprinodon rubrofluviatilis | url=http://txstate.fishesoftexas.org/cyprinodon%20rubrofluviatilis.htm | publisher=Texas Freshwater Fishes, Texas State University - San Marcos, Department of Biology | access-date=26 August 2019 }} 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 {{cvt|15|to|-1.8|C|0}} in less than 24 hours (in the coldest temperatures they bury into the substrate).{{cite journal| author1=Bennett, W.A. | author2=T.L. Beitinger | year=1997 | title=Temperature Tolerance of the Sheepshead Minnow, Cyprinodon variegatus | journal=Copeia | volume=1997 | issue=1 | pages=77–87 | doi=10.2307/1447842| jstor=1447842 }} Some populations of C. nevadensis tolerate water temperatures between {{cvt|2|and(-)|44|C|F|0}}, and C. salinus live in waters where the temperature may change by as much as {{cvt|19|C-change}} in a day and {{cvt|40|C-change}} in a season.{{cite web| author1=Moyle, P.B. | author2=R.M. Yoshiyame | author3=J.E. Williams | author4=E.D. Wirkamanayake | year=1995 | title=Fish Species of Special Concern in California | url=https://dfgsecure.dfg.ca.gov/wildlife/nongame/publications/docs/fish_ssc.pdf | publisher=California Department of Fish and Game | url-status=dead | archive-url=https://web.archive.org/web/20120323024900/https://dfgsecure.dfg.ca.gov/wildlife/nongame/publications/docs/fish_ssc.pdf | archive-date=23 March 2012 }} 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.

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.{{cite iucn |author=NatureServe |date=2013 |title=Cyprinodon arcuatus |volume=2013 |page=e.T202375A15362268 |doi=10.2305/IUCN.UK.2013-1.RLTS.T202375A15362268.en |access-date=12 November 2021}}{{cite iucn |author=Contreras-Balderas, S. |author2=Almada-Villela, P. |author3=Palmer-Newton, A. |date=2019 |title=Cyprinodon ceciliae |volume=2019 |page=e.T6177A3107534 |doi=10.2305/IUCN.UK.2019-2.RLTS.T6177A3107534.en |access-date=12 November 2021}}{{cite iucn |author=Schmitter-Soto, J. |date=2019 |title=Cyprinodon inmemoriam |volume=2019 |page=e.T6176A3107502 |doi=10.2305/IUCN.UK.2019-2.RLTS.T6176A3107502.en |access-date=12 November 2021}}{{cite iucn|url=https://www.iucnredlist.org/details/6168/0 |title=Cyprinodon spp. |author1=Contreras-Balderas, S.|author2=Almada-Villela, P.|year=1996 |access-date=1 March 2013}} C. arcuatus was restricted to springs in the US state of Arizona and probably also in the Mexican state of Sonora, while all the others were restricted to spring systems in Mexico.{{cite journal| author1=Contreras-MacBeath, T. | author2=M.B. Rodríguez | author3=V. Sorani | author4=C. Goldspink| author5=G.M. Reid | year=2014 | title=Richness and endemism of the freshwater fishes of Mexico | journal=Journal of Threatened Taxa | volume=6 | issue=2 | pages=5421–5433 | doi=10.11609/JoTT.o3633.5421-33 | doi-access=free }}{{cite book | editor1=Ceballos, G. | editor2=E.D. Pardo | editor3=L.M. Estévez | editor4=H.E. Pérez | year=2016 | title=Los peces dulceacuícolas de México en peligro de extinción | publisher=Fondo de Cultura Económic | isbn=978-607-16-4087-1 }}

Three species from southwestern Nuevo León, C. alvarezi, C. longidorsalis and C. veronicae, have become extinct in the wild, only surviving in captivity.{{cite iucn |author=Valdes Gonzales, A. |date=2019 |title=Cyprinodon alvarezi |volume=2019 |page=e.T6144A3104750 |doi=10.2305/IUCN.UK.2019-2.RLTS.T6144A3104750.en |access-date=12 November 2021}}{{cite iucn |author=Valdes Gonzales, A. |date=2019 |title=Cyprinodon longidorsalis |volume=2019 |page=e.T6174A3107266 |doi=10.2305/IUCN.UK.2019-2.RLTS.T6174A3107266.en |access-date=12 November 2021}} 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.{{cite journal| author1=Martin, C.H. | author2=Crawford, J.E. | author3=Turner, B.J. | author4=Simons, L.H. | year=2016 | title=Diabolical survival in Death Valley: recent pupfish colonization, gene flow and genetic assimilation in the smallest species range on earth | journal=Proc Biol Sci | volume=283 | issue=1823 | doi=10.1098/rspb.2015.2334 | pmc=4795021 | pmid=26817777 | page=20152334}} Several others have very small remaining populations in the wild. 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.{{cite web|url=http://www.fws.gov/nevada/protected_species/fish/species/dhp/dhp.html|title=Devils Hole Pupfish|publisher=U.S. Fish and Wildlife Service|date=December 2, 2013|access-date=7 October 2018}} Primary threats to pupfish are habitat loss due to water extraction, drought and pollution, and introduced species.

Behavior

=Feeding=

File:Cyprinodon Desquamator.jpg, 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.{{cite journal| author1=Martin, C. | author2=P.C. Wainwright | year=2011 | title=Trophic novelty is linked to exceptional rates of morphological diversification in two adaptive radiations of Cyprinodon pupfish | journal=Evolution | volume=65 | issue=8 | pages=2197–2212 | doi=10.1111/j.1558-5646.2011.01294.x| pmid=21790569 | doi-access=free }} Some species mainly feed on small animals like aquatic insects. C. variegatus, a species that otherwise has a diet typical of pupfish, will clean other fish by feeding on parasites on their body.{{cite web| author1=Hassan-Williams, C. | author2=T.H. Bonner | year=2007 | title=Cyprinodon variegatus | url=http://txstate.fishesoftexas.org/cyprinodon%20variegatus.htm | publisher=Texas Freshwater Fishes, Texas State University - San Marcos, Department of Biology | access-date=26 August 2019 }}

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).

=Breeding=

{{multiple image | direction = horizontal | image1 = Macularius stimulation.jpg | image2 = Salt Creek pupfish.jpg | width1 = 239 | footer = 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, although some may reach as much as three years. 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. 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 {{cvt|2|and(-)|44|C|F|0}}, they only breed from {{cvt|24|to(-)|30|C|F|0}}. However, there are exceptions like C. rubrofluviatilis that will breed at a relative wide range from {{cvt|13|to(-)|34|C|F|0}}. 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. When breeding, males assume a relatively bright nuptial coloration.

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.{{cite journal| author1=Leiser, J.K. | author2=M. Itzkowitz | year=2002 | title=The relative costs and benefits of territorial defense and the two conditional male mating tactics in the Comanche Springs pupfish (Cyprinodon elegans) | journal=Acta Ethol | volume=5 | issue=1 | pages=65–72 | doi=10.1007/s10211-002-0066-1 | s2cid=42642403 }} In at least some species, small males will attempt to fertilize eggs by sneaking into a territory of a larger male. 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. 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. The eggs of Cyprinodon pupfish are adhesive and stick to the substrate, or they are covered in sand.

Species

File:Cyprinodon brontotheroides.jpg (shown) and C. desquamator are both restricted to hypersaline lakes in the Bahamas]]

File:Sonoyta pupfish.jpg in shallow water, which is preferred by most species in this genus]]

File:Julimes pupfish (Cyprinodon julimes) male.jpg was only scientifically described in 2009]]

File:Cyprinodon rubrofluviatilis.jpg, a relatively common species from Texas]]

File:Cyprinodon desquamator.jpg

File:Cyprinodon macrolepis.tif

There are currently 49 recognized species in this genus:

References

{{Reflist}}

{{Taxonbar|from=Q1726419}}

{{Authority control}}

Category:Cyprinodontidae

Category:Ovoviviparous fish

Category:Freshwater fish genera

Category:Taxa named by Bernard Germain de Lacépède