''Cyprinodon'' is a genus of pupfishes found in waters that range from fresh to hypersaline. The genus is primarily found in Mexico, the

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and southern United States (

Arizona Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...

California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...

Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...

Nevada Nevada ( ; ) is a landlocked state in the Western United States. It borders Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. Nevada is the seventh-most extensive, th ...

New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...

Oklahoma Oklahoma ( ; Choctaw language, Choctaw: , ) is a landlocked U.S. state, state in the South Central United States, South Central region of the United States. It borders Texas to the south and west, Kansas to the north, Missouri to the northea ...

and

Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...

), but ''C. variegatus'' occurs as far north as

Massachusetts Massachusetts ( ; ), officially the Commonwealth of Massachusetts, is a U.S. state, state in the New England region of the Northeastern United States. It borders the Atlantic Ocean and the Gulf of Maine to its east, Connecticut and Rhode ...

and along the entire

Gulf of Mexico The Gulf of Mexico () is an oceanic basin and a marginal sea of the Atlantic Ocean, mostly surrounded by the North American continent. It is bounded on the northeast, north, and northwest by the Gulf Coast of the United States; on the southw ...

coastline, and ''C. dearborni '' and ''C. variegatus'' are found in northern South America.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. ''Cyprinodon'' are small; the largest reaches 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 The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

, and saw a rapid evolutionary radiation afterwards. The only known fossil species from the genus is '' C. breviradius'' from the

late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

or early

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Death Valley National Park Death Valley National Park is a national park of the United States that straddles the California–Nevada border, east of the Sierra Nevada. The park boundaries include Death Valley, the northern section of Panamint Valley, the southern sect ...

(originally considered part of the

Late Eocene The Priabonian is, in the International Commission on Stratigraphy, ICS's geologic timescale, the latest age (geology), age or the upper stage (stratigraphy), stage of the Eocene epoch (geology), Epoch or series (stratigraphy), Series. It spans ...

-aged Titus Canyon Formation).

Distribution and habitat

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 A Mexican State (), officially the Free and Sovereign State (), is a constituent federative entity of Mexico according to the Constitution of Mexico. Currently there are 31 states, each with its own constitution, government, state governor, a ...

U.S. state In the United States, a state is a constituent political entity, of which there are 50. Bound together in a political union, each state holds governmental jurisdiction over a separate and defined geographic territory where it shares its so ...

Hispaniola Hispaniola (, also ) is an island between Geography of Cuba, Cuba and Geography of Puerto Rico, Puerto Rico in the Greater Antilles of the Caribbean. Hispaniola is the most populous island in the West Indies, and the second-largest by List of C ...

, 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 and the latter to an shelf in a spring pool, but ''C. longidorsalis'' is now only found in captivity as its habitat has disappeared. 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 Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

in Lake Chichancanab in

Quintana Roo Quintana Roo, officially the Free and Sovereign State of Quintana Roo, is one of the 31 states which, along with Mexico City, constitute the 32 administrative divisions of Mexico, federal entities of Mexico. It is divided into municipalities of ...

, Mexico, and three (the endemic ''C. brontotheroides'' and ''C. desquamator'', and the widespread ''C. variegatus'') live in the

hypersaline lake A hypersaline lake is a landlocked body of water that contains significant concentrations of sodium chloride, brines, and other salts, with saline levels surpassing those of ocean water (3.5%, i.e. ). Specific microbial species can thrive i ...

s 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 Salinity () is the saltiness or amount of salt (chemistry), salt dissolved in a body of water, called saline water (see also soil salinity). It is usually measured in g/L or g/kg (grams of salt per liter/kilogram of water; the latter is dimensio ...

and temperature range is very broad, like those living in

hot spring A hot spring, hydrothermal spring, or geothermal spring is a Spring (hydrology), spring produced by the emergence of Geothermal activity, geothermally heated groundwater onto the surface of the Earth. The groundwater is heated either by shallow ...

s (taken to the extreme in ''C. julimes'' in water up to , and ''C. pachycephalus'' up to ), and those of hypersaline habitats where the salinity far exceeds that of sea water. 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 in less than 24 hours (in the coldest temperatures they bury into the substrate). Some populations of ''C. nevadensis'' tolerate water temperatures between , and ''C. salinus'' live in waters where the temperature may change by as much as in a day and in a season. 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 In taxonomy, an undescribed taxon is a taxon (for example, a species) that has been discovered, but not yet formally described and named. The various Nomenclature Codes specify the requirements for a new taxon to be validly described and named. U ...

popularly known as the " Perrito de Sandia" are already extinct. ''C. arcuatus'' was restricted to springs in the US state of

and probably also in the Mexican state of

Sonora Sonora (), officially Estado Libre y Soberano de Sonora (), is one of the 31 states which, along with Mexico City, comprise the Administrative divisions of Mexico, Federal Entities of Mexico. The state is divided into Municipalities of Sonora, 72 ...

, while all the others were restricted to spring systems in Mexico. Three species from southwestern Nuevo León, ''C. alvarezi'', ''C. longidorsalis'' and ''C. veronicae'', have become extinct in the wild, only surviving in captivity. A few other Mexican species still considered

endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...

or vulnerable by the

IUCN The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...

, including at least ''C. maya'', ''C. simus'' and ''C. verecundus'' of Lake Chichancanab, also appear to only survive in captivity. 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

; in recent decades its population has fluctuated between a few tens and a few hundred individuals. Primary threats to pupfish are habitat loss due to water extraction, drought and pollution, and introduced species.

Behavior

Feeding

Most ''Cyprinodon'' species feed on

algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...

cyanobacteria Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...

and

detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...

, but may also supplement their diet with small crustaceans and aquatic insect larvae. 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. In the two places where several species live together they have diverged into different niches, including the fish-eating ''C. maya'' (Lake Chichancanab),

zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...

-eating ''C. simus'' (Lake Chichancanab), amphipod- and

bivalve Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...

-eating ''C. labiosus'' and ''C. verecundus'' (Lake Chichancanab), scale-eating ''C. desquamator'' (San Salvador Island lakes), and

ostracod Ostracods, or ostracodes, are a Class (biology), class of the crustacean, Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 33,000 species (only 13,000 of which are extant taxon, extant) have been identified,Brandão, S.N.; Antoni ...

- and

gastropod Gastropods (), commonly known as slugs and snails, belong to a large Taxonomy (biology), taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and fro ...

-eating ''C. brontotheroides'' (San Salvador Island lakes).

Breeding

''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 , they only breed from . However, there are exceptions like ''C. rubrofluviatilis'' that will breed at a relative wide range from . 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 A territory is an area of land, sea, or space, belonging or connected to a particular country, person, or animal. In international politics, a territory is usually a geographic area which has not been granted the powers of self-government, ...

and displays to visiting females that will lay their eggs inside the territory. 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

There are currently 49 recognized species in this genus: * '' Cyprinodon albivelis'' W. L. Minckley & R. R. Miller, 2002 (Whitefin pupfish) * '' Cyprinodon alvarezi'' R. R. Miller, 1976 (Potosi pupfish) * †'' Cyprinodon arcuatus'' W. L. Minckley & R. R. Miller, 2002 (Santa Cruz pupfish) * '' Cyprinodon artifrons'' C. L. Hubbs, 1936 (Yucatán pupfish) * '' Cyprinodon atrorus'' R. R. Miller, 1968 (Bolson pupfish) * '' Cyprinodon beltrani'' Álvarez, 1949 (Blackfin pupfish) * '' Cyprinodon bifasciatus'' R. R. Miller, 1968 (Cuatro Cienegas pupfish) * '' Cyprinodon bobmilleri'' Lozano-Vilano & Contreras-Balderas, 1999 (San Ignacio pupfish) * '' Cyprinodon bondi'' G. S. Myers, 1935 (Hispaniola pupfish) * '' Cyprinodon bovinus'' S. F. Baird & Girard, 1853 (Leon Springs pupfish) * †'' Cyprinodon breviradius'' R. R. Miller, 1945 (

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

;

Early Pliocene Early may refer to: Places in the United States * Early, Iowa, a city * Early, Texas, a city * Early Branch, a stream in Missouri * Early County, Georgia * Fort Early, Georgia, an early 19th century fort Music * Early B, stage name of Jamaican d ...

of California) * '' Cyprinodon brontotheroides'' C. H. Martin & Wainwright, 2013 (Durophage pupfish) * †'' Cyprinodon ceciliae'' Lozano-Vilano & Contreras-Balderas, 1993 (Villa Lopez pupfish) * '' Cyprinodon dearborni'' Meek, 1909 * '' Cyprinodon desquamator'' C. H. Martin & Wainwright, 2013 (Scale-eating pupfish) * '' Cyprinodon diabolis''

Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...

, 1930 (Devil's Hole pupfish) * '' Cyprinodon elegans'' S. F. Baird & Girard, 1853 (Comanche Springs pupfish) * '' Cyprinodon eremus'' R. R. Miller & Fuiman, 1987 (Sonoyta pupfish) * '' Cyprinodon esconditus'' Strecker, 2002 (Hidden pupfish) * '' Cyprinodon eximius'' Girard, 1859 (Conchos pupfish) * '' Cyprinodon fontinalis'' M. L. Smith & R. R. Miller, 1980 (Carbonera pupfish) * '' Cyprinodon higuey'' C. M. Rodriguez & M. L. Smith, 1990 (Black-and-blue Pupfish) * '' Cyprinodon hubbsi'' Carr, 1936 (Lake Eustis minnow) * †'' Cyprinodon inmemoriam'' Lozano-Vilano & Contreras-Balderas, 1993 (Charco Azul pupfish) * '' Cyprinodon julimes'' De la Maza-Benignos & Vela-Valladares, 2009 (Julimes pupfish) * '' Cyprinodon labiosus''

Humphries Humphries is a surname, and may refer to: * Barry Humphries (1934–2023), Australian comedian, creator of characters Dame Edna Everage and Sir Les Patterson * Carla Humphries (born 1988), American-born Filipina actress and commercial model, also ...

& R. R. Miller, 1981 (Thicklip pupfish) * '' Cyprinodon laciniatus'' C. L. Hubbs & R. R. Miller, 1942 (Bahama pupfish) * '' Cyprinodon latifasciatus'' Garman, 1881 (Parras pupfish) * ''

Cyprinodon longidorsalis ''Cyprinodon longidorsalis'', the cachorrito de charco palmal or La Palma pupfish, is a species of fish in the family Cyprinodontidae. It was Endemism, endemic to the Ojo de Agua la Presa in southwestern Nuevo Leon state in Mexico, but became ext ...

'' Lozano-Vilano & Contreras-Balderas, 1993 (La Palma pupfish) * '' Cyprinodon macrolepis'' R. R. Miller, 1976 (Largescale pupfish) * '' Cyprinodon macularius'' S. F. Baird & Girard, 1853 (Desert pupfish) * '' Cyprinodon maya''

& R. R. Miller, 1981 (Maya pupfish) * '' Cyprinodon meeki'' R. R. Miller, 1976 (Mezquital pupfish) * '' Cyprinodon nazas'' R. R. Miller, 1976 (Nazas pupfish) * '' Cyprinodon nevadensis'' C. H. Eigenmann & R. S. Eigenmann, 1889 ** '' Cyprinodon nevadensis amargosae'' R. R. Miller, 1948 (Amargosa River pupfish) ** †'' Cyprinodon nevadensis calidae'' R. R. Miller, 1948 (Tecopa pupfish) ** '' Cyprinodon nevadensis mionectes'' R. R. Miller, 1948 (Ash Meadows pupfish) ** '' Cyprinodon nevadensis nevadensis'' C. H. Eigenmann & R. S. Eigenmann, 1889 (Amargosa pupfish) ** '' Cyprinodon nevadensis pectoralis'' R. R. Miller, 1948 (Warm Springs pupfish) ** '' Cyprinodon nevadensis shoshone'' R. R. Miller, 1948 (Shoshone pupfish) * '' Cyprinodon nichollsi'' M. L. Smith, 1989 (Jaragua pupfish) * '' Cyprinodon pachycephalus'' W. L. Minckley & C. O. Minckley, 1986 (Bighead pupfish) * '' Cyprinodon pecosensis'' A. A. Echelle & A. F. Echelle, 1978 (Pecos pupfish) * '' Cyprinodon pisteri'' R. R. Miller & W. L. Minckley, 2002 (Palomas pupfish) * '' Cyprinodon radiosus'' R. R. Miller, 1948 (Owens pupfish) * '' Cyprinodon riverendi'' ( Poey, 1860) (synonym: ''Cyprinodon jamaicensis'', Fowler, 1939) * '' Cyprinodon rubrofluviatilis'' Fowler, 1916 (Red River pupfish) * '' Cyprinodon salinus'' R. R. Miller, 1943 ** '' Cyprinodon salinus milleri'' LaBounty &

Deacon A deacon is a member of the diaconate, an office in Christian churches that is generally associated with service of some kind, but which varies among theological and denominational traditions. Major Christian denominations, such as the Cathol ...

, 1972 (Cottonball marsh pupfish) ** '' Cyprinodon salinus salinus'' R. R. Miller, 1943 (Death Valley pupfish, Salt Creek pupfish) * '' Cyprinodon salvadori'' Lozano-Vilano, 2002 (Bocochi pupfish) * '' Cyprinodon simus''

& R. R. Miller, 1981 (Boxer pupfish) * '' Cyprinodon suavium'' Strecker, 2005 * '' Cyprinodon tularosa'' R. R. Miller & A. A. Echelle, 1975 (White Sands pupfish) * '' Cyprinodon variegatus'' (Lacépède, 1803) ** '' Cyprinodon variegatus baconi'' Breder, 1932 ** '' Cyprinodon variegatus ovinus'' ( Mitchill, 1815) ** '' Cyprinodon variegatus variegatus'' ( Lacépède, 1803) (Sheepshead minnow) * '' Cyprinodon verecundus''

, 1984 (Largefin pupfish) * '' Cyprinodon veronicae'' Lozano-Vilano & Contreras-Balderas, 1993 (Charco Palma pupfish)

References

{{Authority control Cyprinodontidae Ovoviviparous fish Freshwater fish genera Taxa named by Bernard Germain de Lacépède