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{{Short description|Genus of fishes}}
{{Automatic taxobox
{{Automatic taxobox
| image = Owens pupfish (Cyprinodon radiosus).jpg
| image = Owens pupfish (Cyprinodon radiosus).jpg
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| taxon = Cyprinodon
| taxon = Cyprinodon
| authority = [[Bernard Germain de Lacépède|Lacépède]], 1803
| authority = [[Bernard Germain de Lacépède|Lacépède]], 1803
| type_species = ''Cyprinodon variegatus''
| type_species = ''[[Cyprinodon variegatus]]''
| type_species_authority = Lacépède, 1803
| type_species_authority = Lacépède, 1803
| synonyms =
| synonyms =
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}}
}}


'''''Cyprinodon''''' is a genus of small [[pupfish]]es found in fresh, [[Brackish water|brackish]] and salt water. 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.<ref name=FishBase>{{FishBase genus | genus = Cyprinodon| month = October | year = 2018}}</ref><ref name=Martin2013>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.</ref> Many species have tiny ranges and are highly threatened, in some cases already extinct.<ref name=Martin2013/><ref name=Balderas1996>{{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 }}</ref> ''Cyprinodon'' are small; the largest reaches {{convert|10|cm|abbr=on}} in length and most other species only reach about half that.<ref name=FishBase/>
'''''Cyprinodon''''' is a genus of [[pupfish]]es found in waters that range from [[Fresh water|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.<ref name=FishBase>{{FishBase genus | genus = Cyprinodon| month = October | year = 2018}}</ref><ref name=Martin2013>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.</ref> Many species have tiny ranges and are highly threatened, in some cases already extinct.<ref name=Martin2013/><ref name=Balderas1996>{{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 }}</ref><ref>{{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}}</ref> ''Cyprinodon'' are small; the largest reaches {{convert|10|cm|abbr=on}} in length and most other species only reach about half that size.<ref name=FishBase/>


==Distribution and habitat==
==Distribution and habitat==
[[File:Cyprinodon variegatus (S0103) (12598764423).jpg|thumb|''[[Sheepshead minnow|Cyprinodon variegatus]]'' is the most widespread member of the genus.]]
[[File:Cyprinodon diabolis Wales, 1930.jpg|thumb|''[[Cyprinodon diabolis]]'' is very rare; its entire native range is the [[Devils Hole]] in Nevada]]
[[File:Cyprinodon diabolis Wales, 1930.jpg|thumb|''[[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 [[The Bahamas|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.<ref name=Balderas1996/><ref name=Helfman2009>{{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 | page=413 | publisher=Wiley-Blackwell | isbn=978-1-4051-2494-2 }}</ref> While most ''Cyprinodon'' species have separate distributions, seven (''C. beltrani'', ''C. esconditus'', ''C. labiosus'', ''C. maya'', ''C. simus'', ''C. suavium'' and ''C. verecundus'') are [[Endemism|endemic]] in [[Lake Chichancanab]] in [[Quintana Roo]], Mexico,<ref>Contreras Balderas, S., and W. Bussing |(2013). ''[http://www.feow.org/ecoregions/details/175 Yucatan]''. Freshwater Ecoregions of the World. Retrieved 1 March 2013.</ref><ref name=Strecker2006>{{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 }}</ref> 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.<ref name=Martin2013/> In a few other cases separate species do come into contact towards the edges of their distributions where they often [[Hybrid (biology)|hybridize]], notably ''C. eximius'' X ''C. pachycephalus '' and ''C. atrorus'' X ''C. bifasciatus'', but also ''C. variegatus'' in places where it has been [[Introduced species|introduced]] by humans into the ranges of other ''Cyprinodon'' species.<ref name=Martin2013/>
A few ''Cyprinodon'' species have quite large ranges, notably ''[[Sheepshead minnow|C. variegatus]]'', but the vast majority have small ranges, typically restricted to one or two [[Mexican states]] or [[U.S. state]]s, [[Hispaniola]], or a [[The Bahamas|Bahaman]] island. ''[[Cyprinodon longidorsalis|C. longidorsalis]]'' and ''[[Devils Hole pupfish|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.<ref name=Balderas1996/><ref name=Helfman2009>{{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 }}</ref> While most ''Cyprinodon'' species have separate distributions, seven (''C. beltrani'', ''C. esconditus'', ''C. labiosus'', ''C. maya'', ''C. simus'', ''C. suavium'' and ''C. verecundus'') are [[Endemism|endemic]] in [[Lake Chichancanab]] in [[Quintana Roo]], Mexico,<ref>Contreras Balderas, S., and W. Bussing |(2013). ''[http://www.feow.org/ecoregions/details/175 Yucatan]''. Freshwater Ecoregions of the World. Retrieved 1 March 2013.</ref><ref name=Strecker2006>{{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 }}</ref> and three (the endemic ''C. brontotheroides'' and ''C. desquamator'', and the widespread ''C. variegatus'') live in the [[hypersaline lake]]s on [[San Salvador Island]] in the Bahamas.<ref name=Martin2013/> In a few other cases separate species do come into contact towards the edges of their distributions where they often [[Hybrid (biology)|hybridize]], notably ''C. eximius'' X ''C. pachycephalus '' and ''C. atrorus'' X ''C. bifasciatus'', but also ''C. variegatus'' in places where it has been [[Introduced species|introduced]] by humans into the ranges of other ''Cyprinodon'' species.<ref name=Martin2013/>


Although the individual ''Cyprinodon'' species often have a highly specific habitat, overall the genus occurs in a remarkably 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 spring]]s (taken to the extreme in ''C. julimes'' in water up to {{cvt|46|C|disp=or}})<ref name=Clarke2017>{{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 }}</ref> and those of hypersaline habitats where the salinity far exceeds that of sea water.<ref name=Otto1973>{{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 }}</ref> Certain species may even experience very large variations in the temperature and salinity over a relatively short period, like some populations of ''C. variegatus'' where the water 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), and certain species in the [[Death Valley]] region where the water temperature may change by as much as {{cvt|19|C-change}} in a day and {{cvt|40|C-change}} in a season,<ref name=Bennett1997>{{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 }}</ref> and the salinity may vary from less than one-third of that of sea water to almost five times as much as sea water in a season.<ref name=Otto1973/>
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 spring]]s (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}}),<ref name=Clarke2017>{{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 }}</ref><ref>{{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 }}</ref> and those of hypersaline habitats where the salinity far exceeds that of sea water.<ref name=Otto1973>{{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 }}</ref><ref name=Williams2007rubrofluviatilis>{{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 }}</ref> 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).<ref name=Bennett1997>{{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 }}</ref> 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.<ref name=Bennett1997/><ref name=Moyle1995>{{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 }}</ref> 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.<ref name=Otto1973/>


==Conservation status==
==Conservation status==
Most species in the genus are seriously threatened. ''C. arcuatus'', ''C. ceciliae'', ''C. inmemoriam'', ''C. latifasciatus'' and an [[undescribed species]] popularly known as the "[[Perrito de Sandia]]" are already extinct,<ref name=IUCNarcuatus>{{IUCN|id=202375 |title=''Cyprinodon arcuatus'' |assessors=Smith, K. & Darwall, W.R.T. |version=2018.1 |year=2013 |accessdate=8 October 2018}}</ref><ref>{{IUCN|id=6177 |title=''Cyprinodon ceciliae'' |assessors=Contreras-Balderas, S. & Almada-Villela, P. |version=2012.2 |year=1996 |accessdate=1 March 2013}}</ref><ref>{{IUCN|id=6176 |title=''Cyprinodon inmemoriam'' |assessors=Contreras-Balderas, S. & Almada-Villela, P. |version=2012.2 |year=1996 |accessdate=1 March 2013}}</ref><ref>{{IUCN|id=6155 |title=''Cyprinodon latifasciatus'' |assessors=Contreras-Balderas, S. & Almada-Villela, P. |version=2012.2 |year=1996 |accessdate=1 March 2013}}</ref><ref>{{IUCN|id=6168 |title=''Cyprinodon spp.'' |assessors=Contreras-Balderas, S. & Almada-Villela, P. |version=2012.2 |year=1996 |accessdate=1 March 2013}}</ref> while ''C. alvarezi'', ''C. longidorsalis'' and ''C. veronicae'' have become [[extinct in the wild]], only surviving in captivity.<ref>{{IUCN|id=6144 |title=''Cyprinodon alvarezi'' |assessors=Contreras-Balderas, S. & Almada-Villela, P. |version=2012.2 |year=1996 |accessdate=1 March 2013}}</ref><ref>{{IUCN|id=6174 |title=''Cyprinodon longidorsalis'' |assessors=Contreras-Balderas, S. & Almada-Villela, P. |version=2012.2 |year=1996 |accessdate=1 March 2013}}</ref><ref name=MacBeath2014>{{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 | url=https://threatenedtaxa.org/index.php/JoTT/article/view/966/1728 | journal=Journal of Threatened Taxa | volume=6 | issue=2 | pages=5421–5433 | doi=10.11609/JoTT.o3633.5421-33 }}</ref> ''C. arcuatus'' was restricted to springs in the American state of [[Arizona]] and probably also in the Mexican state of [[Sonora]],<ref name=IUCNarcuatus/> while all the others were restricted to spring systems in Mexico, with ''C. latifasciatus'' in southern [[Coahuila]] and the remaining in southwestern [[Nuevo León]].<ref name=MacBeath2014/><ref name=Ceballos2016>{{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 | isbn=978-607-16-4087-1 }}</ref> A few other Mexican species still considered [[Endangered species|endangered]] by the [[IUCN]], including at least ''C. maya'', ''C. simus'' and ''C. verecundus'' of Lake Chichancanab, also appear to only survive in captivity.<ref name=Martin2011/><ref name=Martin2016>{{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 }}</ref> Several others have very small remaining populations in the wild.<ref name=Ceballos2016/> 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 fluctuation between a few tens and a few hundred individuals.<ref name=FWSdiabolis>{{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|accessdate=7 October 2018}}</ref> Primary threats to pupfish are habitat loss due to water extraction, drought and pollution, and introduced species.<ref name=Balderas1996/><ref name=Ceballos2016/>
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.<ref name=Moyle1995/><ref name=IUCNarcuatus>{{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}}</ref><ref>{{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}}</ref><ref>{{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}}</ref><ref>{{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}}</ref> ''C. arcuatus'' was restricted to springs in the US state of [[Arizona]] and probably also in the Mexican state of [[Sonora]],<ref name=IUCNarcuatus/> while all the others were restricted to spring systems in Mexico.<ref name=MacBeath2014>{{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 }}</ref><ref name=Ceballos2016>{{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 }}</ref>

Three species from southwestern Nuevo León, ''C. alvarezi'', ''C. longidorsalis'' and ''C. veronicae'', have become [[extinct in the wild]], only surviving in captivity.<ref name=MacBeath2014/><ref>{{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}}</ref><ref>{{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}}</ref> A few other Mexican species still considered [[Endangered species|endangered]] or [[Vulnerable species|vulnerable]] by the [[IUCN]], including at least ''C. maya'', ''C. simus'' and ''C. verecundus'' of Lake Chichancanab, also appear to only survive in captivity.<ref name=Martin2011/><ref name=Martin2016>{{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}}</ref> Several others have very small remaining populations in the wild.<ref name=Ceballos2016/> 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.<ref name=FWSdiabolis>{{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}}</ref> Primary threats to pupfish are habitat loss due to water extraction, drought and pollution, and introduced species.<ref name=Balderas1996/><ref name=Ceballos2016/>


==Behavior==
==Behavior==
===Feeding===
[[File:Cyprinodon Desquamator.jpg|thumb|''[[Cyprinodon desquamator]]'', the only known [[scale-eating]] species of pupfish]]
[[File:Cyprinodon Desquamator.jpg|thumb|''[[Cyprinodon desquamator]]'', the only known [[scale-eating]] species of pupfish]]


''Cyprinodon'' pupfish typically feed on [[algae]] and [[detritus]], but in the two places where several species live together they have diverged into different [[Ecological niche|niches]], including the [[Piscivore|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).<ref name=Martin2013/><ref name=Martin2011>{{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}}</ref>
Most ''Cyprinodon'' species feed on [[algae]], [[cyanobacteria]] and [[detritus]], but may also supplement their diet with small crustaceans and aquatic insect larvae.<ref name=Moyle1995/><ref name=Martin2011>{{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 }}</ref> Some species mainly feed on small animals like aquatic insects.<ref name=Williams2007rubrofluviatilis/> ''C. variegatus'', a species that otherwise has a diet typical of pupfish, will [[Cleaner fish|clean other fish]] by feeding on parasites on their body.<ref name=Williams2007variegatus>{{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 }}</ref>

In the two places where several species live together they have diverged into different [[Ecological niche|niches]], including the [[Piscivore|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).<ref name=Martin2013/><ref name=Martin2011/>

===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).<br/>
⚫
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,<ref name=Moyle1995/> although some may reach as much as three years.<ref name=Williams2007variegatus/> 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.<ref name=Moyle1995/> 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}}.<ref name=Moyle1995/> However, there are exceptions like ''C. rubrofluviatilis'' that will breed at a relative wide range from {{cvt|13|to(-)|34|C|F|0}}.<ref name=Williams2007rubrofluviatilis/> 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.<ref name=Moyle1995/> When breeding, males assume a relatively bright nuptial coloration.<ref name=Moyle1995/>

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 (animal)|territory]] and displays to visiting females that will lay their eggs inside the territory.<ref name=Moyle1995/><ref name=Leiser2002>{{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 }}</ref> In at least some species, small males will attempt to fertilize eggs by [[Sneaking (biology)|sneaking]] into a territory of a larger male.<ref name=Leiser2002/> 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.<ref name=Moyle1995/> 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.<ref name=Moyle1995/> The eggs of ''Cyprinodon'' pupfish are adhesive and stick to the substrate,<ref name=Moyle1995/> or they are covered in sand.<ref name=Williams2007variegatus/>


==Species==
==Species==
[[File:Cyprinodon brontotheroides.jpg|thumb|''[[Cyprinodon brontotheroides]]'' (shown) and ''C. desquamator'' are both restricted to [[hypersaline]] lakes in the Bahamas]]
[[File:Cyprinodon brontotheroides.jpg|thumb|''[[Cyprinodon brontotheroides]]'' (shown) and ''C. desquamator'' are both restricted to [[hypersaline]] lakes in the Bahamas]]
[[File:Macularius stimulation.jpg|thumb|''[[Cyprinodon macularius]]'', pair during courtship (male above)]]
[[File:Sonoyta pupfish.jpg|thumb|''[[Cyprinodon eremus]]'' in shallow water, which is preferred by most species in this genus]]
[[File:Julimes pupfish (Cyprinodon julimes) male.jpg|thumb|''[[Cyprinodon julimes]]'' was only scientifically described in 2009]]
[[File:Cyprinodon rubrofluviatilis.jpg|thumb|''[[Cyprinodon rubrofluviatilis]]'', a relatively common species from Texas]]
[[File:Cyprinodon rubrofluviatilis.jpg|thumb|''[[Cyprinodon rubrofluviatilis]]'', a relatively common species from Texas]]
[[File:Cyprinodon desquamator.jpg|alt=scale-eating pupfish in the wild|thumb|Cyprinodon desquamator (shown), the scale-eating pupfish, in its natural habitat on San Salvador Island, Bahamas]]
⚫
[[File:Salt Creek pupfish.jpg|thumb|''[[Cyprinodon salinus]]'', spawning pair (male with yellowish back)]]
[[File:Cyprinodon macrolepis.tif|thumb|Cyprinodon macrolepis male in an aquarium.]]

There are currently 49 recognized species in this genus:<ref name=FishBase/><ref name=Martin2013/>
There are currently 49 recognized species in this genus:<ref name=FishBase/><ref name=Martin2013/>
* ''[[Cyprinodon albivelis]]'' <small>[[Wendell L. Minckley|W. L. Minckley]] & [[Robert Rush Miller|R. R. Miller]], 2002</small> (Whitefin pupfish)
* ''[[Cyprinodon albivelis]]'' <small>[[Wendell L. Minckley|W. L. Minckley]] & [[Robert Rush Miller|R. R. Miller]], 2002</small> (Whitefin pupfish)
Line 57: Line 73:
* ''[[Cyprinodon eximius]]'' <small>[[Charles Frédéric Girard|Girard]], 1859</small> (Conchos pupfish)
* ''[[Cyprinodon eximius]]'' <small>[[Charles Frédéric Girard|Girard]], 1859</small> (Conchos pupfish)
* ''[[Cyprinodon fontinalis]]'' <small>[[Michael Leonard Smith|M. L. Smith]] & [[Robert Rush Miller|R. R. Miller]], 1980</small> (Carbonera pupfish)
* ''[[Cyprinodon fontinalis]]'' <small>[[Michael Leonard Smith|M. L. Smith]] & [[Robert Rush Miller|R. R. Miller]], 1980</small> (Carbonera pupfish)
* ''[[Cyprinodon higuey]]'' <small>[[Carlos Manuel Rodriguez|C. M. Rodriguez]] & [[Michael Leonard Smith|M. L. Smith]], 1990</small> (Black-and-blue Pupfish)
* ''[[Cyprinodon higuey]]'' <small>[[Carlos Manuel Rodriguez|C. M. Rodriguez]] & [[Michael Leonard Smith|M. L. Smith]], 1990</small> (Black-and-blue Pupfish)<ref>{{cite iucn |author=Lyons, T.J. |date=2021 |title=''Cyprinodon higuey'' |volume=2021 |page=e.T82958669A82962221 |doi=10.2305/IUCN.UK.2021-1.RLTS.T82958669A82962221.en |access-date=12 November 2021}}</ref>
* ''[[Cyprinodon hubbsi]]'' <small>[[Archie Carr|Carr]], 1936</small> (Lake Eustis minnow)
* ''[[Cyprinodon hubbsi]]'' <small>[[Archie Carr|Carr]], 1936</small> (Lake Eustis minnow)
* [[Extinction|†]]''[[Cyprinodon inmemoriam]]'' <small>[[María de Lourdes Lozano-Vilano|Lozano-Vilano]] & [[Salvador Contreras-Balderas|Contreras-Balderas]], 1993</small> (Charco Azul pupfish)
* [[Extinction|†]]''[[Cyprinodon inmemoriam]]'' <small>[[María de Lourdes Lozano-Vilano|Lozano-Vilano]] & [[Salvador Contreras-Balderas|Contreras-Balderas]], 1993</small> (Charco Azul pupfish)
Line 63: Line 79:
* ''[[Cyprinodon labiosus]]'' <small>[[Julian M. Humphries|Humphries]] & [[Robert Rush Miller|R. R. Miller]], 1981</small> (Thicklip pupfish)
* ''[[Cyprinodon labiosus]]'' <small>[[Julian M. Humphries|Humphries]] & [[Robert Rush Miller|R. R. Miller]], 1981</small> (Thicklip pupfish)
* ''[[Cyprinodon laciniatus]]'' <small>[[Carl Leavitt Hubbs|C. L. Hubbs]] & [[Robert Rush Miller|R. R. Miller]], 1942</small> (Bahama pupfish)
* ''[[Cyprinodon laciniatus]]'' <small>[[Carl Leavitt Hubbs|C. L. Hubbs]] & [[Robert Rush Miller|R. R. Miller]], 1942</small> (Bahama pupfish)
* [[Extinction|†]]''[[Cyprinodon latifasciatus]]'' <small>[[Samuel Garman|Garman]], 1881</small> (Parras pupfish)
* ''[[Cyprinodon latifasciatus]]'' <small>[[Samuel Garman|Garman]], 1881</small> (Parras pupfish) <ref name=Gonzales2019>{{cite iucn |author=Valdes Gonzales, A. |date=2019 |title=''Cyprinodon latifasciatus'' |volume=2019 |page=e.T6155A3105057 |doi=10.2305/IUCN.UK.2019-2.RLTS.T6155A3105057.en |access-date=12 November 2021}}</ref>
* ''[[Cyprinodon longidorsalis]]'' <small>[[María de Lourdes Lozano-Vilano|Lozano-Vilano]] & [[Salvador Contreras-Balderas|Contreras-Balderas]], 1993</small> (La Palma pupfish)
* ''[[Cyprinodon longidorsalis]]'' <small>[[María de Lourdes Lozano-Vilano|Lozano-Vilano]] & [[Salvador Contreras-Balderas|Contreras-Balderas]], 1993</small> (La Palma pupfish)
* ''[[Cyprinodon macrolepis]]'' <small>[[Robert Rush Miller|R. R. Miller]], 1976</small> (Largescale pupfish)
* ''[[Cyprinodon macrolepis]]'' <small>[[Robert Rush Miller|R. R. Miller]], 1976</small> (Largescale pupfish)
Line 72: Line 88:
* ''[[Cyprinodon nevadensis]]'' <small>C. H. Eigenmann & R. S. Eigenmann, 1889</small>
* ''[[Cyprinodon nevadensis]]'' <small>C. H. Eigenmann & R. S. Eigenmann, 1889</small>
** ''[[Cyprinodon nevadensis amargosae]]'' <small>[[Robert Rush Miller|R. R. Miller]], 1948</small> (Amargosa River pupfish)
** ''[[Cyprinodon nevadensis amargosae]]'' <small>[[Robert Rush Miller|R. R. Miller]], 1948</small> (Amargosa River pupfish)
** ''[[Cyprinodon nevadensis calidae]]'' <small>[[Robert Rush Miller|R. R. Miller]], 1948</small> (Tecopa pupfish)
** [[Extinction|†]]''[[Cyprinodon nevadensis calidae]]'' <small>[[Robert Rush Miller|R. R. Miller]], 1948</small> (Tecopa pupfish)
** ''[[Cyprinodon nevadensis mionectes]]'' <small>[[Robert Rush Miller|R. R. Miller]], 1948</small> (Ash Meadows pupfish)
** ''[[Cyprinodon nevadensis mionectes]]'' <small>[[Robert Rush Miller|R. R. Miller]], 1948</small> (Ash Meadows pupfish)
** ''[[Cyprinodon nevadensis nevadensis]]'' <small>[[Carl H. Eigenmann|C. H. Eigenmann]] & [[Rosa Smith Eigenmann|R. S. Eigenmann]], 1889</small> (Amargosa pupfish)
** ''[[Cyprinodon nevadensis nevadensis]]'' <small>[[Carl H. Eigenmann|C. H. Eigenmann]] & [[Rosa Smith Eigenmann|R. S. Eigenmann]], 1889</small> (Amargosa pupfish)
Line 102: Line 118:


{{Taxonbar|from=Q1726419}}
{{Taxonbar|from=Q1726419}}
{{Authority control}}


[[Category:Cyprinodon| ]]
[[Category:Cyprinodon| ]]

Latest revision as of 17:44, 29 April 2024

Cyprinodon
Cyprinodon radiosus
Scientific classification Edit this classification
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
  • 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.[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

[edit]
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

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 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

[edit]

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

[edit]

Feeding

[edit]
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

[edit]
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

[edit]
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
Cyprinodon julimes was only scientifically described in 2009
Cyprinodon rubrofluviatilis, a relatively common species from Texas
scale-eating pupfish in the wild
Cyprinodon desquamator (shown), the scale-eating pupfish, in its natural habitat on San Salvador Island, Bahamas
Cyprinodon macrolepis male in an aquarium.

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

References

[edit]
  1. ^ a b c Froese, Rainer; Pauly, Daniel (eds.). "Species in genus Cyprinodon". FishBase. October 2018 version.
  2. ^ a b c d e f g h 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. ^ a b c 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. ^ a b 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. ^ a b c 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. ^ a b 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. ^ a b c d e f g h i j k l 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. ^ a b 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. ^ a b 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. ^ a b c 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. ^ a b c 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. ^ a b c 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. ^ a b 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.
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